VOLUME 31 OCTOBER 1880

CALIFORNIA COOPERATIVE OCEANIC INVESTIGATIONS

VOLUME 31 January 1 to December 31,1989

Cooperating Agencies:

CALIFORNIA DEPARTMENT OF AND GAME UNIVERSITY OF CALIFORNIA, SCRIPPS INSTITUTION OF NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION, NATIONAL MARINE FISHERIES SERVICE CALCOFI COORDINATOR Patricia Wolf

EDITOR Julie Olfe

SPANISH EDITOR Carina Lange

This report is not copyrighted, except where otherwise indicated, and may be reproduced in other publications provided credit is given to the California Cooperative Oceanic Fisheries Investigations and to the author(s). Inquiries concerning this report should be addressed to CalCOFl Coordinator, Cali- fornia Department of Fish and Game, 330 Golden Shore, Suite 50, long Beach, CA 90802.

EDITORIAL BOARD lzadore Barrett Richard Klingbeil Michael Mullin

Published October 7990, la Jolla, California ISSN 0575-3377 CalCOFl Rep., Vol. 31,1990

CONTENTS

In Memoriam ...... 5 I. Reports, Review, and Publications Report of the CalCOFI Committee ...... , ...... 7 Review of Some California Fisheries for 1989 ...... 9 Publications ...... 22 11. Fortieth Anniversary Symposium of the CalCOFI Conference, 1989 OCEAN OUTLOOK: GLOBAL CHANGE AND THE MARINE ENVIRONMENT ...... 25 Presentation to Roger Revelle ...... 28 Remarks by the Honorable Leo McCarthy, Lieutenant Governor of California ...... , ...... 30 Remarks by the Honorable John Knauss, Under Secretary of Commerce for Oceans and Atmosphere ...... 33 Panel Discussion: What Society Needs from Ocean Scientists in Preparing for Global Change. Robert Sulnick, By von She6 Edward Frieman, Bert Larkins, John McGowan, Boyce Thorne Millev, and Havry Scheiber ...... , ...... , ...... 37 111. Scientific Contributions California Marine Research and the Founding of Modern Fisheries Oceanography: CalCOFI’s Early Years, 1947-1964. Havry N. Scheibev ...... 63 Settlement ofJuvenile California Halibut, Paralichthys cal~ovnicus,along the Coasts of Los Angeles, Orange, and San Diego Counties in 1989. M. James Allen and Kevin ?: Herbinson . .. 84 Genetic Structure of White Seabass Populations from the Southern California Bight Region: Applications to Hatchery Enhancement. Devin M. Bartley and Donald B. Kent ...... 97 The Vertical Distribution and Feeding Habits of Two Common Midwater (Letrroglossus stilbius and Stenobvachius leucopsavus) off Santa Barbara. Gvegov M. Cailliet and AlfYed W Ebeling ...... , ...... 106 The Market for Fish Meal and Oil in the United States: 1960-1988 and Future Prospects. Cynthia]. Thomson ...... , ...... 124 Bathymetric Patterns in Size, Age, Sexual Maturity, Water Content, and Caloric Density of

Dover Sole, Microstomus -.pacificus. John R. Huntev, john L. Butlev, Carol Kimbrell, and Eric A. Lynn ...... 132 Monitoring Interannual Changes in Spawning Area of Pacific Sardine (Savdinopssagax). Paul E. Smith ...... 145 Instructions to Authors ...... 152 CalCOFI Basic Station Plan ...... inside back cover

Indexed in Curvent Contents. Abstracted in Aquatic Sciences and Fishevies Abstracts and Oceanic Abstracts.

IN MEMORIAM CalCOFl Rep., Vol. 31,1990

IN MEMORIAM

James R. Thrailkill 1921- 1990

Jim Thrailkill (left) receives his forty-year service pin from Reuben Lasker.

CalCOFI lost a friend when Jim Thrailkill died in in India during World War I1 and a stint as a surveyor Boise, Idaho, after a brief illness at the age of 68. for the U. S. Geological Survey. After retirement from the National Marine Fisheries His long career in fisheries began in 1949 when he Service in April 1986, Jim returned to his native transferred to the sardine investigation of the then Boise where he could enjoy the good trout U.S. Fish arid Wildlife Service in San Diego. He and beautiful surroundings of nearby streams. joined others to begin the federal government’s co- At the time of his retirement, Jim was the leader operative investigations with Scripps Institution of of the Coastal and Pacific Fisheries Investigation Oceanography into the reasons for the catastrophic within the Coastal Fisheries Resources Division at decline in Pacific sardine landings. His first cruise the Southwest Fisheries Center. He received his was on the Block Doii

5 IN MEMORIAM CalCOFl Rep., Vol. 31,1990

he supervised the conversion of field and laboratory Jim was a wonderfully warm and cheerful man. data procedures to a computerized system. Jim em- He always greeted people with a smile and friendly phasized accuracy and consistency in all aspects of conversation. He never hesitated to offer his help his CalCOFI work and is largely responsible for the when it was needed - and it usually was. Those of high quality of the time series. He was the author us who were fortunate enough to know Jim will of a series of scientific reports on zooplankton always remember his unbounded loyalty, generos- volumes, coauthor of several papers on plankton ity, and kindness. volume loss with time of preservation, and collabo- rator in the development of a high-speed plankton sampler.

6 COMMITTEE REPORT CalCOFl Rep., Vol. 31, 1 990

Part I REPORTS, REVIEW, AND PU BL I CAT10 N S

REPORT OF THE CALCOFI COMMITTEE

The CalCOFI family of member agencies ob- which was incorporated into a stock synthesis esti- served its fortieth anniversary in 1989 by acknowl- mate of anchovy spawning biomass. Unusually low edging past accomplishments and evaluating present water temperatures may have inhibited sexual ma- strengths in preparation for the global challenges of turity, reduced spawning activity, and resulted in a the years to come. The 40-year CalCOFI collection low estimate of spawning biomass, since the 1988 of physical, chemical, biological, and meteorologi- year class appeared large, and total biomass was cal data from the California Current is the most judged to be high. National Marine Fisheries Ser- complete ocean time series in the world, and has led vice (NMFS) and CDFG scientists prepared an to an understanding of the pelagic ecosystem that amendment to the anchovy Management is unmatched in any comparable marine region. Plan to allow a small reduction fishery under cir- CalCOFI has also served as a model of successful cumstances when the total biomass is high but collaboration among diverse agencies. An article in the spawning biomass is below the cutoff level this volume chronicles the early history of CalCOFI for fishing. and the roles of the visionary scientists and managers At the third annual meeting of MEXUS-Pacifico, who partnered in its development. it was suggested that the cooperative scope of this The sardine resource continues to recover, and the fisheries research agreement between the United California Department of Fish and Game (CDFG) States and Mexico be broadened beyond coastal pe- permitted the fifth 1,000-ton quota fishery in as lagic species to include sea lions, sea turtles, and many years. For the second year, the quota, which remote sensing. We continued our routine exchange opened on January 1, 1990, was allocated between of fisheries and biological data, and conducted two northern (200 tons) and southern (800 tons) Califor- cruises in Mexican waters to estimate anchovy egg nia. The 800-ton quota was landed in one week, production. For 1990, we planned a port sampling completing the shortest season to date. workshop, joint egg production cruises to assess an- Fishery-related cruises included a 50-day ground- chovy and sardine biomass, a stock synthesis work- fish survey to collect sablefish eggs and evaluate the shop, and a workshop on fisheries applications of use of the egg production method for determining satellite technology. A workshop on aging pelagic spawning biomass; two cruises to collect sardine fishes was held in Ensenada. eggs off southern California and northern Baja Cal- CalCOFI continued to support the Spanish-Por- ifornia for biomass assessment; two night-lighting tugese Sardine Anchovy Recruitment Program cruises to assess recruitment of juvenile Pacific (SARP). We hosted a meeting to review work ac- mackerel; and one midwater trawl survey to collect complished over the last three years by SARP partic- young anchovy, mackerel, and sardines. The quar- ipants. A planning meeting sponsored by the Inter- terly CalCOFI cruises surveying the southern governmental Oceanographic Commission (IOC) California sector of the California Current were of the United Nations Education, Scientific and completed. In addition, a rapid but intensive hydro- Cultural Organization followed immediately, and graphic survey of the periphery of the station grid included an ad hoc expert consultation session on was conducted just after the summer cruise, and SARP analysis of data from the 1988 biological/physical In a break with tradition, the 1989 CalCOFI survey of the Ensenada Front continued. The pur- Conference was held at the Scripps Institution of chase of a CTD will permit a gradual change in the Oceanography. The symposium, which consisted techniques used on the CalCOFI surveys, improv- of invited addresses and a panel discussion, was or- ing the vertical resolution of data without impairing ganized to honor the fortieth anniversary of the the continuity of the time series. CalCOFI program, and to consider what society We used 1989 CalCOFI collections of anchovy and its policymakers can reasonably expect in terms eggs and larvae to estimate daily egg production, of scientific advice concerning large-scale changes

7 COMMITTEE REPORT CalCOFl Rep., Vol. 31,1990

in the ocean. Other facets of the anniversary celebra- to thank everyone who contributed to volume 31 of tion included the preparation of a brochure and a CalCOFI Reports: editor Julie Olfe for her profes- videotape (for which E. Venrick deserves special sional, thorough work and patient assistance; Span- thanks) describing CalCOFI and some of its notable ish editor Carina Lange; past CalCOFI Coordinator achievements; the construction of two new perma- George Hemingway and current Coordinator Patri- nent exhibits at the Scripps aquarium-museum; a cia Wolf; and the many peer reviewers for their time, presentation to Roger Revelle honoring his role in effort, and suggested improvements to the scientific CalCOFI’s early years; and some unusually elabo- contributions. The reviewers for this volume were rate wining and dining. A special CalCOFI exhibit Alice Alldredge, John Butler, Dan Cohen, Dudley and a continuous showing of the CalCOFI video Chelton, John Cullen, Deborah Day, Thomas Hay- were also part of the NMFS Southwest Fisheries ward, Dennis Hedgecock, Daniel Huppert, Sharon Center twenty-fifth anniversary rededication and Kramer, John Marl-, Milton Love, Alec MacCall, open house. Marc Mangel, Douglas McLain, Geoffrey Moser, Many thanks to the officers and crews who assist Michael Mullin, Tim Parsons, Elizabeth Venrick, us in our work on the University of California RV Robin Waples, and James Waters. New Horizon, the National Oceanic and Atmo- spheric Administration ship David Starr Jordan, the The CalCOFI Committee: Southern California Ocean Studies Consortium RV Izadore Bavvett Yellowfin, the RV Shana Rae, the RV Westwind, and Richard Klingbeil the FVjonathan Michael. The Committee also wishes Michael Mullin

8 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

REVIEW OF SOME CALIFORNIA FISHERIES FOR 1989

California Department of Fish and Game Marine Resources Division 330 Golden Shore, Suite 50 Long Beach, California 90802

Total landings of fishes, crustaceans, echino- the shortfin mako shark and the Pacific angel shark derms, and mollusks decreased slightly (3%) this continued to decline. year, after three years of increase. The 1989 landings Dungeness crab landings showed a slight in- remained well below the ten-year average, by ap- crease, and Pacific Ocean landings continued proximately 20%. However, they exceeded the 1985 to rise for the sixth straight year. Landings of the low by nearly 32%. southern California spiny lobster were the highest Pelagic wetfish landings continued the upward since the mid-1950s. trend that began in 1985, with a 2% gain over last Landings of the California red sea urchin de- year (table 1).Jack mackerel landings nearly doubled creased slightly, and the fishery is likely to come from last year, and market squid landings continued under increasingly restrictive management mea- to be high. Pacific herring landings increased to the sures in 1970. highest level since 1782, and the take of northern Albacore landings declined for the fourth consec- anchovy and Pacific sardine also increased. Pacific utive year and reached an all-time low in 1989. Only mackerel landings decreased by 16%. 10% of the previous 25-year average was landed. Groundfish landings increased slightly, but dif- The sport catch increased, and rockfish retained fered markedly in species composition from last the first-rank position. year. California halibut landings increased slightly but have remained fairly constant over the last nine PACIFIC SARDINE years. The California Department of Fish and Game Landings of swordfish and the common thresher (CDFG) conducted sea surveys in May of 1988 to shark increased in 1989. However, landings for both assess the spawning biomass of the Pacific sardine

TABLE 1 Landings of Pelagic Wetfishes in California (Short Tons) Pacific Northern Pacific Jack Pacific Market Year sardine anchovy mackerel mackerel herring squid Total 1966 439 31,140 2,315 20,431 121 9,513 63,959 1967 74 34,805 583 19,090 136 9,801 64,489 1968 62 15,538 1 567 27,834 179 12,466 57,646 1969 53 67,639 1,179 26,961 85 10,390 106,307 1970 22 1 96,243 31 1 23,873 158 12,295 133,101 1971 149 44,853 78 29,941 120 15,579 90,900 1972 186 69,101 54 25,559 63 10,080 105,043 1973 76 132,636 28 10,308 1,410 6,031 150,489 1974 7 82,691 67 12,729 2,630 14,453 112,577 1975 3 158,510 144 18,390 1,217 11,811 190,075 1976 27 124,919 328 22,274 2,410 10,153 160,l 11 1977 6 111,477 5,975 50,163 5,827 14,122 187,570 1978 5 12,607 12,540 34,456 4,930 18,899 83,437 1979 18 53,881 30,471 18,300 4,693 32,026 129,389 1980 38 47,339 32,645 22,428 8,886 16,957 128,293 1981 31 57,659 42,913 15,673 6,571 25,915 148,762 1982 145 46,364 31,275 29,110 11,322 17,951 136,167 1983 388 4.740 35,882 20,272 8,829 2,001 72,112 1984 259 3,258 46,531 11,768 4,241 622 66,679 1985 653 1,792 38,150 10,318 8,801 11,326 71,040 1986 1,283 2,105 45,503 12,209 8,405 23,454 92,959 1987 2,309 1,595 45,890 13,055 9,258 22,028 94,135 1988 4,172 1,618 47,278 11,379 9,721 41,040 115,208 1989* 4,308 2,700 39,725 21.820 10.134 38.288 116.975 *Preliniinarv

9 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

(Sardinops sugux). The egg production area method mackerel from traditional fishing grounds. The tol- (EPAM) was used to determine if the observed erance limit for incidentally landed sardines mixed spawning area (based on the occurrence of sardine with other fish remained at 35% by weight. eggs) exceeded the minimum critical spawning area Landings from all sources, excluding live bait, of2,300 nautical miles’ (n.mi.>),which is considered totaled 4,308 tons in 1989, as compared to 4,172 tons to indicate a 20,000-ton spawning biomass. A in 1988 and 2,309 tons in 1987 (table 1). Of this year’s 20,000-ton spawning biomass is needed before a di- catch, 93% can be attributed to southern California rected fishery for sardine can be permitted. Results landings, and just under 7% to the northern Califor- indicated spawning activity over an area of about nia allocation of the directed fishery. Sardine sold for 2,508 n.mi.” As a result, on January 1, 1989, a 1,000- approximately $100 per ton. ton directed fishery for sardines was opened for the AB 2351, which became effective in September fourth consecutive year. 1989, allocates the 250-ton dead bait quota among Of the 1,000-ton quota, 800 tons were allocated three geographic regions: 125 tons are reserved for for landings south of Point Buchon, and 200 tons landings south of Point Buchon, 50 tons for landings were allocated for landings to the north. Also estab- north of Point Buchon and south of Pescadero lished were a 350-ton quota (beginning on January Point, and 75 tons for landings north of Pescadero 1) for live bait, and a 250-ton quota (beginning on Point. In addition, all sardine fishing for dead bait March 1) for use as dead bait. purposes must be accompanied by a written order The southern allocation of the 1989 directed fish- from a processor; all fish landed must be kept in a ery closed on January 12, three days earlier than the whole condition; and the receipts must be labeled previous year, with total landings of 924 tons. Of “For Dead Bait Only. ” This is to ensure that no fish these landings, 34% were pure loads and another allocated for dead bait will be used for other pur- 32% contained at least 70% sardines. The fish were poses, and to facilitate monitoring of landings caught by the southern California mackerel purse against the quota. seine fleet and were almost exclusively canned for Biomass estimation cruises using the EPAM were human consumption. again conducted in 1989, and were expanded to in- The northern California directed fishery saw no clude, for the first time, areas off northern Baja Cal- landings until late February, and remained open un- ifornia as far south as Bahia de San Quintin. The til early April. The 23 landings totaled 258 tons, and area off central California north of Point Conception almost all were pure loads of sardines. The fish were was not sampled in 1989 because no spawning activ- all purchased by a single processor and marketed for ity was observed in this area during the 1988 cruise. human consumption. During the 1989 cruise, evidence of spawning was The 250-ton dead bait quota proved difficult to observed along the California coast out to the Chan- monitor. Processors were not required to specifi- nel Islands, from Santa Barbara south to Dana Point, cally declare the intended use of purchased sardines. and offshore in a large area west of Tanner and Cor- Unless a landing exceeded the allowable incidental tez banks (figure l), over a total area of3,280 n.mi.’ tolerance limit (35% sardines by weight, mixed Evidence of spawning off Baja California was lim- with other fish), sardines within that load were gen- ited to a small area totaling 400 n.mi.” As in 1988, erally not declared as dead bait. The quota was the total area over which spawning was observed reached on March 20, when an estimated 250 tons exceeded what is considered to indicate a 20,000-ton had been landed. Live bait landings reported by fish- spawning biomass; as a result, a 1,000-ton directed ermen amounted to 111 tons, with an additional 194 fishery for sardines was scheduled for 1990. tons estimated by Department observers on sport- On December 7, the CDFG held a meeting for fishing partyboats. As in 1988, young-of-the-year members of the sardine industry. The occasion pro- sardines did not make a strong showing in the 1989 vided an opportunity for the CDFG to present the live bait fishery. methods and results of the fishery research that have Incidental landings of sardines in the mackerel led to the current and proposed management direc- fishery totaled 2,876 tons, down 7% from 1988 and tives. The meeting also provided a forum for the reflecting the overall decrease of activity within the industry to express its concerns and intentions re- mackerel fishery. Sardines composed just under 6% garding the sardine resource. of the mackerel landings, up slightly from the 5% observed in 1988. Fishermen continued to complain MARKET SQUID that the abundance of sardines interfered with mack- Market squid (Loligo opulescetzs) landings in 1989 erel fishing, and that sardines are displacing the were 38,288 short tons (table 1): 31,011 tons (81%)

10 FISHERIES REVIEW: 1989 ColCOFl Rep., Vol. 31,1990

125' I I I I I

Crescent Clty

Trlnldad

Eureka

False Cape

Santa Barbara O0 estport u

Fort Brag9 San Nicolas

:HANNEL ISLANDS Pt. Arena

so

Pt. Conceptlon Santa Barbere

LO8 Angel68

Tanner (3 Bank Corter Bank c-.,', ; Sen DleQO -J I I I I I I I I I I 125' 1200

Figure 1. California ports and fishing areas

11 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

were from the southern California fall-winter fish- The Monterey Bay area fishery experienced major ery; 7,274 tons (19%) from northern California changes in 1989. Attracting lights were allowed for (Monterey Bay area) spring-summer fishery; and 3 the first time in many years in the southern bight of tons from other areas. The total ex-vessel value was Monterey Bay. Also, purse seiners were allowed to approximately $5.9 million. Ex-vessel prices typi- fish in this area, which before had been restricted to cally fluctuate from year to year and during the year. the use of lampara nets. By the end of the season, The ex-vessel price paid in southern California con- the Monterey Bay area round haul fleet had all con- tinued to be lower than that paid in the Monterey verted to purse seine gear. area. Southern California fishermen were paid from Despite the widespread use of attracting lights by $120 to $200 per ton, while Monterey’s ex-vessel nearly all the Monterey Bay fleet, the fishing com- price started at $200 per ton and increased to $260 munity appears to be split on the issue of using per ton after a mid-April strike. Within the past 20 lights. Many fishermen contend that lights disrupt years, the ex-vessel price has been as high as $600 squid spawning, which could adversely affect the per ton. fishery in the future. Others point out that southern Most squid were frozen for human consump- California fishermen have used lights for many years tion; some were sold fresh; some were used for dead without any apparent effects on spawning. They bait; and some were used for live bait. Exported also claim that lights allow the fishermen to use shal- squid accounted for a large percentage of the total lower nets, which can be fished off the bottom, thus processed. protecting squid egg clust‘ers attached there. Annual squid landings during the last 20 years In 1987 and 1988, the CDFG authorized three averaged 16,740 tons. Landings during years char- Monterey boats to experiment with purse seines in acterized by El Nifio Southern Oscillation (ENSO), southern Monterey Bay to test their effectiveness such as 1973, 1983, and 1984, were unusually low. relative to lampara nets, and their effects on squid However, large increases in landings in southern egg clusters and bottom habitat. As a result of this California and increased fishing effort in other areas study, all round haul nets were authorized in this area in recent years suggest that the resource has been in 1989. Many fishermen were especially interested underutilized. in the half-purse drum seine, which allows a smaller Before 1986, annual southern California landings crew to work the gear. Continuing low ex-vessel averaged less than one-half of total statewide land- squid prices and difficulties in finding enough crew ings; since 1986, southern California landings have have heightened interest in drum seines. Few boats, risen dramatically, averaging nearly 80% of the to- tal. One reason for southern California’s dramatic rebound following the 1983-84 ENSO is an increase in squid fishing effort around the Channel Islands TABLE 2 off the coast of Santa Barbara (figure 1). During the California Market Squid Landings (Short Tons) 20 years before the 1983-84 ENSO, Santa Barbara Southern State area landings averaged about 1,500 tons per year; Year Monterey California Other __total since then, landings have averaged about 11,700 tons 1970 4,314 7,982 0 12,295 1971 8,323 7,435 trace 15,759 per year. Several large Monterey-based boats now 1972 6,129 3.950 0 10,080 regularly participate in this fishery and deliver to 1973 620 5,140 0 6,031 Santa Barbara area ports. These squid are trucked to 1974 7,248 7,205 0 14,453 1975 2,495 9,316 tracr 11,811 Monterey for processing. 1976 2,511 7,642 0 10,153 During the last 20 years, Monterey Bay area an- 1977 2,234 11,887 1 14,122 nual landings averaged 6,060 tons. Since the 1983- 1978 10,328 8,571 tracr 18,899 1979 14,183 7,842 1 22,026 84 ENSO, annual landings have averaged approxi- 1980 7,856 9,100 1 16,957 mately 5,600 tons, slightly below the 20-year average 1981 14,134 11,779 2 25,915 and well below the annual average of 11,600 tons per 1982 11,670 6,276 5 17,951 1983 542 9-50 509 2,001 year during 1978-82 (table 2). Landings of 7,274 1984 43 1 84 107 622 tons in 1989 made this an above-average year. Of 1985 4,202 7.039 85 11,326 this total, 975 tons were landed at the port of Santa 1986 6,049 16,488 917 23,454 1987 5,269 16,381 378 22,028 Cruz. These squid were caught north of Santa Cruz 1988 5,330 35.348 363 31,040 1989* 7,274 31,011 near Afio Nuevo Island, an area fished only sporad- 3 __38,288 ically during post-ENS0 years. *Prrliminary

12 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

however, switched to drum seines this year, primar- per ton for fish used for pet food. The purse seine ily because a major capital outlay this year was for fleet has seen the mackerel price decline steadily: powerful (and expensive) attracting light systems in the early 1980s the canneries paid $200 per ton, that typically cost’from $16,000 to $20,000. More and over the last few years the price declined to boats will probably convert to drum seines in $155 per ton. the future. By the end of the year, only 15,447 tons of Pacific mackerel had been landed toward the 1989-90 sea- PACIFIC MACKEREL son total. Landings of Pacific mackerel for the year The year began with 19,120 tons of Pacific mack- totaled 39,725 tons (table 1). These are the lowest erel (Scombev japonicits) already landed through the annual landings since 1985 and are 11 YOlower than first half of the 1988-89 season (July 1 through June the previous five-year average. Northern California 30). Current law allows an open fishery when the landings contributed less than 1% to the year’s total. biomass exceeds 150,000 tons. Since the biomass This continues a steady decline in the proportion of was estimated to be 220,000 tons, no quota restric- the catch occurring in Monterey. tions were in effect. In general, market conditions were stable through During the first three months of the year, mack- the year. Processors continued to impose landing erel landings were high, even though the sardine limits, which sometimes were as low as 20 tons per fishery in early January and poor weather through- boat, per day, to limit landings of small fish. Two out the period threatened to interfere with mackerel changes in southern California mackerel processing effort. Large Pacific mackerel, which had been rela- occurred in 1989, Coast Cannery, a pet food canning tively uncommon since the previous October, be- facility operated by Pan Pacific, ceased operation in came more available in March. Landings declined October. In September, Starkist ’ pet food during April, May, and June, as fishermen were oc- production operation was renamed Heinz Pet Prod- casionally hampered by rough weather and some- ucts, after the parent company. times turned their attention to Pacific bonito (Sauda The CDFG continued to conduct “night-light” chiliensis). Catches ofsmall and large fish were made, surveys in which mackerel were sampled by hook although large fish were reportedly difficult to find. and line, in an effort to develop an early, fishery- The 1988-89 season closed on June 30, 1989, with independent index of year class strength. Two a total catch of 43,398 tons of Pacific mackerel. This cruises, one in April and another in May, were con- is only slightly below the previous five-year season ducted this year. Results suggest that the 1988 year average. Although mackerel landings were lower class is at least similar in strength to the 1986 year than in the last two years, revenues for the San Pedro class. Fishery data support this conclusion: the 1988 purse seine fleet from all fish species increased over year class - which as yearlings contributed nearly last year, primarily because of higher landings of 50% by weight of the catch this year, and which bluefin ( Thiinnus thynnits) and record landings appears to be very strong-and the 1986 year of squid. Pacific mackerel contributed 79% to state- class - which contributed 13% -together made up wide landings of mackerel, and nearly 99% of all most of the landings. Pacific mackerel landings were made in southern California. NORTHERN ANCHOVY The 1989-90 season opened on July 1, 1989, with Landings of northern anchovy (Enptdis moudax) no quota restrictions, based on a biomass estimate for reduction purposes in 1989 were limited primar- of 263,000 tons. Landings during the third quarter, ily by poor market conditions. As in the 1987-88 which were only fair, were comparable to the pre- reduction season, high fish meal prices during the vious quarter. Effort was often directed toward 3 988-89 season were not reflected in the price bluefin tuna, and in September, jack mackerel (Za- offered to local fishermen. California processors chuviir symmetuicus) began to dominate landings. thought that an increase in price to $35 per ton Fourth-quarter landings of Pacific mackerel were would attract some fishermen away from other spe- low. Jack mackerel dominated landings in October cies to anchovy, but few fishermen considered the and November, and price disputes between fisher- price to be fair. For this reason, northern processors men and United Food Processors (UFP), a Terminal issued no orders during the latter half of the 1988- Island cannery, inhibited fishing and persisted 89 season. Although processors in the southern re- through the quarter. UFP reportedly wanted to re- gion issued orders for anchovy, local purse seine duce the price paid to fishermen from $135 to $80 fishermen continued to concentrate on more lucra-

13 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1.990

TABLE 3 southern California processors stated that there Anchovy Landings (Short Tons) for Reduction were no plans to reduce anchovies for the rest of the Southern Northern 1989-90 season. No landings were made in the Season area area Total northern area through December 1989, and orders 1967-68 852 5,651 6,503 are not anticipated before the season’s end. 1968-69 25,314 2,736 28,050 Total anchovy landings during 1989 included 120 1969-70 81,453 2,020 83,473 1970-71 80,095 657 tons for reduction, 2,580 tons for nonreduction pur- 1971-72 52,052 1,314 53,366 poses (table l),and 5,064 tons for live bait. Although 1972-73 73,167 2,352 75,519 live bait landings increased from 1988, most live bait 1973-74 109,207 11,380 120,587 1974-75 109,918 6,669 116,587 fishermen rated 1989 as only slightly better than 1975-76 135,619 5,291 140,910 average. 1976-77 101,434 5,007 106,441 1977-78 68,467 7,212 75,679 1978-79 52,696 1,174 53,870 1979-80 33,383 2,365 35,748 PACIFIC HERRING 1980-81 62,161 4,736 66,897 The 1989 annual roe herring catch (Clupeu huven- 1981-82 45,149 4,953 507102 1982-83 4,925 1,270 6,195 gus pullasi) increased 4%, to 10,134 tons (table l), 1983-84 70 1,765 1,835 and the 1988-89 seasonal (December-March) roe 1984-85 78 0 78 herring catch also increased 4%, to 10,022 tons. 1985-86 0 1,595 1,595 California’s 1988-89 roe herring quota of 9,990 tons 1986-87 0 42 42 198 7 - 8 8 0 122 122 was taken, because of a quota overrun of 236 tons in 1988-89* 0 258 258 San Francisco Bay. Even though the 1988-89 To- *Preliminary males Bay roe herring quota was reduced from 750 to 400 tons, there was a quota shortfall of 187 tons in Tomales Bay. In Crescent City Harbor, the 30- tive mackerel and squid. Consequently, no reduc- ton quota was taken; in Humboldt Bay there was a tion landings took place in either the northern or 16-ton shortfall on the 6O-ton quota. southern regions after December. The 1988-89 sea- The 1988-89 San Francisco Bay herring spawn- son closed on June 30, with six landings totaling 258 ing biomass estimate was 66,000 tons; hydroacous- tons (234 MT; table 3). tic and spawn survey estimates agreed. The popu- National Marine Fisheries Service biologists esti- lation declined about 5% from the 1988 estimate mated the 1989 spawning biomass of northern an- because average recruitment of the 1987 year class chovy to be 235,892 tons (214,000 MT) and the total was not strong enough to maintain the increasing biomass to be 1,111,118 tons (1,008,000MT). Nor- trend in abundance apparent since the 1982-83 mally when the spawning biomass is less than El Nifio. 300,000 MT, the Anchovy In Tomales Bay the herring biomass continues to Plan allows for zero take of northern anchovy for decline. The 1988-89 spawning-ground surveys es- reduction purposes. However, spawning biomass timated spawning escapement of only 167 tons. was low because cold sea-surface temperatures Spawning biomass, which includes the catch, was caused one-year-old fish to mature more slowly and only 380 tons. Both are historic lows for Tomales not be actively spawning when the surveys were Bay. The average structure of the Tomales Bay her- conducted. Because of this anomaly, the Pacific Fish- ring catch - primarily age 4- through 7-year-olds - eries Management Council (PFMC) determined appears normal. This does not support the decline that an emergency existed in the northern anchovy in abundance of the Tomales Bay population. It is fishery and requested the secretary of commerce to believed that most of the Tomales Bay population approve regulations that would allow a 5,000-MT is avoiding the bay because of the recent drought reduction fishery during the 1989-90 season. The and lack of freshwater runoff near historic spawn- secretary approved the emergency rule, which went ing areas. into effect on September 25, 1989. Based on 1988-89 spawning biomass estimates, For the first time since the 1984-85 season, reduc- the 1989-90 San Francisco Bay roe herring quota tion landings were recorded in southern California. remained at 9,500 tons. However, because of the lack Two landings totaling 120 tons (109 MT) were de- of spawning escapement in Tomales Bay, further re- livered to a Terminal Island cannery in December strictions were placed on the 1989-90 season. A 1989. The price paid was $40 per ton. However, spawning threshold of 2,000 tons was established for

14 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

the 1989-90 Tomales Bay season, with no fishing month season. This midwater-trawl fishery, located allowed until 2,000 tons of herring had spawned. If off Eureka and Crescent City, has grown from ap- 2,000 tons escapement was reached before January proximately 3,000 MTin 1987 to 7,300 MT in 1989. 31, the fishery would open with a 400-ton quota. In the 1989 Pacific whiting fishery a conflict oc- The 1989-90 season began quickly in San Fran- curred between shore-based and joint venture (JV) cisco Bay, and the December quota of 1,999 tons operations; the shore-based trawlers testified before was taken easily. In the Tomales Bay area no herring the PFMC that the large and highly efficient JV fleet were caught in December, and there was no spawn- dissipated whiting schools within the small operat- ing activity. ing radius of the shore-based fleet. The JV fishery At the beginning of the 1989-90 season herring involves U. S. trawl vessels delivering whiting to buyers were offering $1,000 per ton for gill net her- foreign processing vessels at sea. The fleet concen- ring, the same as last season. However, the price trated its effort off northern California early in the offered for round haul herring dropped to $400 per season. Although CDFG scientists and others could ton. Japanese buyers are willing to pay more for the not verify this alleged JV impact, it was cited by larger, better-quality herring caught by gill nets. processors as the cause for failure to meet a domestic 1989 production goal of 15,000 MT. A reduced whiting optimum yield (OY)and an increased JV GROU NDFlSH demand in 1990 are expected to intensify conflicts California’s 1989 commercial groundfish harvest within the industry. was 40,510 MT, with an ex-vessel value of approxi- Despite limits on trip poundage and frequency for mately $28,879,000. All-species 1989 landings in- deepwater-assemblage landings of sablefish (Ano- creased only 3%, or 1,090 MT, from 1988 but plopoma jimbvia), Dover sole, arrowtooth flounder differed markedly in species composition (table 4). (Athevesthes stomias), and thornyheads, a robust Setnet landings continued their recent decline, con- Asian market drove thornyhead landings to a record tributing less than 5% of all 1989 groundfish land- high of 5,319 MT. Much of the 70% increase in ings. The general historical pattern of landings by thornyhead landings since 1987 is due to the devel- gear continued during 1989. Bottom and midwater opment of a market for headed-and-gutted long- trawl landings accounted for 86.2% of total land- spine thornyheads (Sebastolobus altivelis), which are ings, followed by line gear (6.6%), setnets (4.8%), typically too small to fillet for domestic consump- and traps (2.4%). The size and composition of the tion. Additional regulations and reduced demand trawl, trap, and longline fleet did not differ mark- apparently contributed to a 6% drop from 1988 in edly from recent years. Rockfish (Sebastes spp.), Dover sole landings. Dover sole (Micvostomus paci_fictis), Pacific whiting Federal and state regulations for 1989 affected the (Merlucciusproductus), and thornyheads (Sebastolobiis harvest of sablefish, Dover sole, thornyheads, and spp.) were the principal species harvested in 1989. widow rockfish (Sebastes entomelns). A coastwide, The domestic shore-based Pacific whiting fishery Washington-Oregon-California (WOC), widow in California achieved record landings during its six- rockfish OY of 12,400 MT (300 MT greater than in 1988), with a 30,000 pound-per-week trip limit was imposed. Excellent fishing conditions in the first TABLE 4 quarter of 1989 accelerated widow rockfish land- California Groundfish Landings (Metric Tons) ings. As a result, the PFMC’s Groundfish Manage-

~ - ~- - Percent ment Team projected that a 51 Yo reduction in rate of Species 1988 1989 change landings would be required to extend the fishery to -~__ -~ Dover sole 8,176 7,713 -6 year’s end. Consequently, the PFMC reduced the English sole 1,062 1,015 -4 trip limit to 10,000 pounds per week, or 20,000 Petrale sole 785 840 7 pounds biweekly, effective April 26, 1989. In early Rex sole 840 735 - 13 Thoriiyheads 4,524 5,319 17 October a by-catch-only trip limit of 3,000 pounds Widow rockfish 1,817 1,566 - 15 was imposed to further slow the fishery. The widow Other rockfish 9,846 9,978 1 rockfish quota was eventually filled, and the fishery Lingcod 873 1,262 45 Sablefish 3,784 3,583 -2 was closed in mid-December. The midseason re- Pacific whiting 6,541 7,302 12 strictions undoubtedly contributed to the 15% Other groundfish 1,142 1.197 5 -~ ~~ ~ ~~ reduction in California’s widow rockfish catch; fish- Total 39,420 40,510 3 ~~~~ ~~ ermen reported that fishable aggregations were

15 FISHERIES REVIEW: 3989 CalCOFl Rep., Vol. 31,1990

available and quite vulnerable to trawl gear during (figure 1) were 5.42, 0.90, 1.28, and 0.24 million most of the year off northern California. pounds, respectively. Production in Crescent City Regulation of the WOC sablefish fishery in- almost doubled, and total northcoast landings ex- creased in complexity during 1989. The year began ceeded 1987-88 seasonal landings by 2.22 million with an OY range of 10,400 to 11,000 MT. The pounds. A total of 318 vessels made 5,436 trips and intent was to manage toward 10,400 MT, with a averaged 1,399 pounds per trip. The price paid to 600-MT reserve to accommodate by-catch and gear the on December 1, opening day, was other than trawl if OY was attained before year’s $1.25 per pound. end. Initial allocations, after a set-aside of 22 MT for The 1988-89 season in the San Francisco/Bodega Native Americans, were 5,397 MT (52%) for trawl Bay area ended with a landing total of 1.44 million and 4,981 MT (48%) for other gear. To maintain a pounds. This is slightly less than half of the total for year-round trawl fishery, a trip limit was imposed of the previous season. Approximately 215 boats made 1,000 pounds, or 45% of the deepwater assemblage a total of 2,866 trips for an average of 501 pounds of sablefish, Dover sole, thornyheads, and arrow- per trip. tooth flounder - whichever was greater. The intent was to discourage targeting on sablefish, while al- lowing sablefish landings from the deepwater as- PACIFIC OCEAN SHRIMP semblage fishery. However, this trip limit did not Statewide landings of Pacific Ocean shrimp (Pun- slow landings sufficiently. At its April 1989 meet- dulus jovduni) in 1989 increased to 13.3 million ing, the PFMC modified the trip limit and trans- pounds, from 11.1 million pounds landed in 1988. ferred the 600-MT reserve and 400-MT other-gear This was the second largest total ever and the sixth quota to the trawl fishery in order to minimize trawl consecutive rise in statewide landings. Areas of pro- discards late in the year. The revised trip limit re- duction were Area A (Oregon border to False Cape) stricted the deepwater assemblage to one landing Area B-1 (False Cape to Point Arena), and Area C per week of not more than 30,000 pounds total. Of (Pigeon Point to the Mexican border; figure 1). this total, sablefish could constitute 1,000 pounds or Shrimp landings at Area A ports totaled 12.5 mil- 25% by weight, whichever was greater. On Octo- lion pounds - a 2.2 million pound increase over 1988 ber 4, PFMC removed the assemblage restrictions deliveries. These landings tie with 1978 as the sec- but retained the sablefish limits until year’s end. Pre- ond largest ever. The total landings comprised 11.74 liminary analysis revealed that the assemblage trip- million pounds from Area A waters, 250,000 frequency and percentage restrictions effectively pounds from Area B-1, and 465,000 pounds from prevented premature quota attainment. California Oregon waters. The season opened on April 1, with trawl sablefish landings of 2,200 MT accounted for fishermen receiving a split price of $0.40 per pound approximately 40% of WOC landings. for shrimp at or below 140 per pound and $0.25 per Directed nontrawl sablefish fishing was termi- pound for smaller shrimp. Shrimpers were on strike nated on July 17, when only 200 MT of quota re- for 26 days in July over a price disagreement, which mained for incidental catches. A 100-pound trip was finally settled at $0.35 per pound for the larger limit subsequently was imposed and remained in size group (140 count or better), with no guarantee effect until early October. In response to testimony of any payment for smaller shrimp. from the Newport, California, dory fleet and north- A total of 56 boats (36 single-rigged and 20 dou- ern California rockfish longline representatives, ble-rigged) delivered shrimp to Area A ports during PFMC increased the limit to 2,000 pounds per trip, 1989, down one boat from 1988. Single-rigged ves- or 20% of all groundfish aboard. California ac- counted for 1,383 MT, or 31%, of the 4,500 MT sels had an average seasonal catch rate of 543 pounds landed by nontrawl gears. per hour, an increase of 55 pounds per hour over 1988. Double-riggers averaged 842 pounds per hour, up from 758 pounds per hour in 1988. DUNGENESS CRAB One-year-old shrimp made up 65% of the catch California Dungeness crab (Cuncev rnugistev) land- in April and 85% in October during 1989. This ings during the 1988-89 season totaled 9.2 million was approximately a 10% decrease in one-year-olds pounds, only slightly more than the 1987-88 land- from 1988. There were no incoming year-class ings of 8.7 million pounds. (zero-aged) shrimp found in the sampled catch, Landings for the northern California ports of the first year-class failure since the total fishery Crescent City, Trinidad, Eureka, and Fort Bragg failure in 1983.

16 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

Landings in area B-1 were 833,000 pounds this only 422 fish, making 1989 one the poorest harpoon season, compared to 379,000 pounds in 1988. Four years on record. Drift gillnetters reported land- local single-rigged boats, along with one double- ing 11,190 fish, nearly the same number reported rigged and two single-rigged vessels from Crescent in 1988. City and one single-rigged boat from Santa Cruz, Fish taken early in the season (August-Septem- fished the B-1 area this year. Ninety-one percent of ber) were much larger (averaging 150 pounds) than the catch was landed at Noyo Harbor in Fort Bragg. fish caught later in the year (averaging 125 pounds). Forty percent of the Fort Bragg landings occurred Catch locations were centered off San Francisco, in April and 73% in April, May, and June combined. Morro Bay, and San Diego (figure 1). One local boat accounted for 54% of the Fort Bragg CPUE for gill net boats remained nearly identical landings. to 1988. Both years had a catch rate of two fish per Catch-per-unit-of-effort (CPUE), measured as day, per boat. The CPUE for harpoon gear was 0.33 pounds landed per delivery, started at 14,600 pounds fish per day, per boat in 1989 and 0.40 fish per day, per delivery in April. CPUE declined to 9,000 pounds per boat in 1988. in May and June, climbed back to 14,200 pounds in Common thresher shark (Alopias vulpincrs) land- August, and declined again to 4,000 pounds in ings in California during 1989 reached 647,865 October. One-year-olds made up 27% of the sam- pounds, 17% more than the 550,000 pounds in 1988. pled catch in April, when ovigerous females were No thresher sharks were landed in Oregon or Wash- also noted. In May, one-year-olds composed 73% of ington during 1989 because these states ended their the sampled catch and ranged from 40% to 52% of permit fishery, mainly because of decreased interest the catch through the end of the season. There were by local fishermen, higher incidental catch of marine no zero-aged shrimp noted during the season. The mammals and leatherback turtles, and concern for count per pound increased in May, causing several the status of the resource. Currently, the Pacific fishers to return to groundfish until the States Marine Fisheries Commission is coordinating count improved. the development of a coastwide management plan. The total shrimp catch for Area C in 1989 was Shortfin mako shark (Iruvus oxyvinchus) landings 24,000 pounds, down considerably from the decreased 20% from 1988. Of the 388,312 pounds 380,000 pounds landed in 1988. This was the least taken, 46% was caught by the experimental drift productive year since 1978, when no shrimp were longline fleet and the remainder by drift gill net landed in Area C. Four single-rigged vessels made boats. Most of the fish were taken off central and five trips and averaged 314 pounds per hour. southern California during the summer and fall. The California Fish and Game Commission re- authorized the use of drift longline gear for 1990 SWORDFISH AND SHARK under the conditions of a 175,000-pound quota Landings of swordfish (Xiphius gladitis) for 1989 and the development of a market for blue shark rose to 2.8 million pounds, a 15% increase from (Pvionace glaucu), specifying that 40,000 pounds be 1988 (table 5). Harpoon fishermen reported landing marketed for human consumption. Pacific angel shark (Sqciatina calgovnica) landings TABLE 5 in 1989 continued their decline for the third straight Landings of Selected Shark Species and Swordfish (Pounds) year, reaching only 267,577 pounds. A management plan establishing a minimum size limit was enacted Shortfin Common Pacific in 1989 to protect juveniles and a portion of the Year mako shark Swordfish thresher shark angel shark spawning stock. Reduced availability, decreased 1977 19,911 51 1,388 129,522 366 1978 26,765 2,604,233 302,054 82,383 market demand, and the size limit appear responsi- 1979 35,079 586,579 735,726 128,295 ble for the lower landings. The fishery continued to 1980 154,529 1,197,187 1,805,978 110,037 be centered off Santa Barbara and Ventura counties, 1981 274,217 1,142,897 1,973,412 268,640 1982 527,006 1,677,020 2,396.960 317,953 although some landings occurred in San Diego and 1983 322,854 2,601,600 1,722,056 351,344 San Pedro (figure 1). 1984 239,687 4,429,540 1,662,587 632,937 1985 225,535 5,196,685 1,540,770 1,237.810 1986 473,608 3,845,932 606,583 1,241,130 CALIFORNIA HALIBUT 1987 602,718 2,741,015 525,076 940,187 1988 488,136 2,484,428 549,516 487,278 California halibut (Pavulichthyr culfovtzicus) land- 1989* 388.312 2.85O. 734 647.865 267.577 ings for 1989 were 550 MT, 9% more than the 505 *Preliminary MT recorded in 1988 (table 6). Catches over the last

17 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

five years have remained fairly constant, averaging Only 303 permits were issued, down 5.6% from the 542 MT, with landings for 1989 exceeding the 13- 1987-88 level of 321. The number of permittees has year average of 448 MT. During 1989, 55% of the decreased each season since 1984-85. halibut landings occurred north of Point Concep- The remaining fishermen, however, may be fish- tion. Landings south of Point Conception accounted ing a greater number of traps, keeping effort high. for the remaining 45%, a 7% decrease from 1988. Most of the catch was taken early in the season: 50% The highest landings occurred during winter and in October, 19% in November, and 12% in Decem- fall, with peak catches in February (58 MT) and ber, The remaining 19% was caught from January October (60 MT). Entangling nets (trammel and set to March. gill nets) accounted for 51% of all halibut taken, Ex-vessel price ranged from $5.00 to $7.00 per followed by trawl (20%), unknown gear (l6%), pound, averaging about $5.50. With landings at hook and line (1 1YO), and other gears (2%). Most of 650,000 pounds, the fishery was worth $3.6 million the trawl-caught (91 Yo) and hook-and-line-caught to the fishermen, a $1.1 million (43%) increase over (93%) halibut were taken off central California. The the previous season. southern California area accounted for nearly 61 '/a of all halibut taken by entangling nets; 39% came ALBACORE from north of Point Conception. Ex-vessel prices In 1989, albacore (Thcrnnus alulungu) landings in for California halibut ranged from $0.45 per pound California reached an all-time low. Only 914 tons of in San Francisco to $5.35 per pound in Ventura, and albacore were brought in; this is approximately 10% averaged $2.25 per pound statewide. of the previous 25-year average. For the past five years, commercial catches of albacore have declined CALIFORNIA SPINY LOBSTER substantially. From 1984 to 1987, season totals de- The southern California spiny lobster (Punulinrs creased by 50% each year. In 1988 and 1989, the rate interruptus) fishery landed 650,000 pounds during of decline slowed to 20% per year (figure 2). Fishing the 1988-89 season (first Wednesday in October effort this year was moderate, with 225 boats partic- to first Wednesday after March 15), making it the ipating in the fishery. Of these, 78 landed over a ton best season since 1955-56, when 790,000 pounds of albacore. were landed. The season had a promising start in July. Catches Historically, landings of lobster from California were reported at Rosa Bank and Geronimo Island, waters peaked at 1.1 million pounds in the 1949-50 off southern Baja California, as well as off the cen- season. Seasonal landings generally declined over tral California coast. In addition, toward the end of the next 25 years, reaching a low of 152,000 pounds July, a good sport fishery developed off southern in 1974-75. Since then, there has been a general California. As the season progressed, however, it upward trend. became apparent that the center of fishing activity The 1988-89 season's total is an increase of was once again off the Washington coast. Seventy 173,000 pounds (36%) from the 1987-88 season, in percent of California's albacore fleet spent August spite of a slight decrease in participating fishermen. through September in the north. A few vessels even followed the albacore as far north as the Queen TABLE 6 California Halibut Landings (Metric Tons)

~~~~ ~ North of South of "1 Year Pt. Conception Pt. Conception Total ~~ ~ ~~ ~- 1977 25 186 21 1 1978 34 165 199 1979 54 205 259 1980 90 23 1 3'71 1981 163 409 572 1982 206 339 545 1983 256 248 504 1984 153 345 198 1985 144 429 573 1986 240 312 552 0 ....,. ..,...... , .... 1987 192 347 530 1960 1965 1970 1975 1980 1985 1990 1988 229 276 505 1989* 305 245 550 Year - __ ~~ ~~ ~ ~ ~~ ~ *Preliminary Figure 2. Annual albacore landings for California, 1960-89

18 FISHERIES REVIEW: 1989 CalCOFl Rep.,Vol. 31,1990

Charlotte Islands, Canada, before returning home TABLE 7 in October. Drift gill net boats fishing primarily off Ridgeback Prawn and Spot Prawn Landings (1,000s of central California had incidental catches of four al- Pounds) bacore per day throughout the season. The fish Year Ridgeback prawn Spot prawn caught off California ranged between 11 and 15 1980 276 69 pounds on the average. Very few large fish (greater 1981 193 369 than 25 pounds) were seen. 1982 141 300 1983 157 109 Price agreements between the Western Fishboat 1984 623 49 Owners Association and Pan Pacific and Starkist 1983 905 64 canneries began at $1,500 per ton for fish over 9 1986 672 102 1987 242 88 pounds and $1,000 per ton for fish 9 pounds and 1988 143 167 under. This was a drop of $200 per ton from the 1989* 176 179 1988 price and was due mainly to the large quantities *Preliminary of albacore coming in from Japan. In addition, ves- sels landing at ports other than Terminal Island were Originally caught in traps, spot prawns were pre- charged a shipping fee, which reduced the price paid dominantly caught by trawl by the mid-1970s. With by another $200 per ton. By the end of August, the the recent increasing demand for live products, trap- shipping fee was eliminated to encourage fishermen ping is on the increase. For the second year, log data to sell their loads to the canneries instead of directly indicate that just over half the catch is taken by traps. to the public. Overall, dockside sales increased 3% Spot prawns may be trawled by permit from Feb- in 1989. ruary 1 through October 31. During the restricted Oceanic and market conditions did not appear to period, an incidental catch of 50 pounds is allowed. influence the poor season in California this year. A Spot prawns can be harvested by trap year-round. strong corridor (warm water/cold water front) de- Log data from 11 boats showed a healthy CPUE of veloped in June and should have led albacore into 75 pounds per hour. Trawling took place in the Santa southern California waters and then along the coast. Barbara Channel, Santa Monica Bay, and off Santa The season was poor, so albacore fishermen Catalina Island (figure 1). Trapping occurred in the were not drawn away to that fishery. The poor same locations and also off San Diego. Ex-vessel albacore season in California can generally be price in the Santa Barbara region was $3.50 to $5.00 attributed to a lack of large albacore off southern per pound. California and an above-average season off Wash- ington. Fishermen had a different opinion: many considered foreign high-seas drift gillnetting to be SEA URCHIN the cause. In 1989 the red sea urchin (Stvongylocentvotusfvun- ciscunus) fishery continued to be one of the major fisheries in the state. Landings for 1989 are estimated RIDGEBACK AND SPOT PRAWN to be 50.9 million pounds, a 2.1% decrease from Ridgeback prawn ( Sicyoniu ingentis) are fished 1988 (table 8). Northern California landings are commercially, primarily by otter trawl. They may down 12.4% from 1988, whereas those from south- be trawled by permit from October 1 through May ern California increased 12.6%. Once again, Fort 31. During the restricted period an incidental catch Bragg led all ports, with 30% of the statewide total. of 50 pounds is allowed. Landings for 1989 were The southern California ports of Santa Barbara, approximately 176,104 pounds, 23% greater than Ventura-Oxnard, and San Pedro-Los Angeles had the previous year’s catch (table 7). Most of the catch 11’7’0, l6%, and 15% of the statewide total. The came from the Santa Barbara Channel (figure 1). reduction in northern California landings is also re- Log data showed a CPUE of 66 pounds per hour, flected by a 20% drop in average pounds per landing virtually unchanged from last year. The average ex- from 1988. These decreases are attributed to the vessel price in the Santa Barbara region was $1.25 continued reduction of high-density, virgin stocks per pound. in northern California. Spot prawn (Pundalus plutycevos) landings in- Divers, using surface-supplied air, harvest sea ur- creased to 179,718 pounds in 1989, about 8% more chins by raking them into mesh bags, which are than in 1988 (table 7). The spot prawn is a larger then air lifted to the surface and winched aboard the shrimp and brings a higher price than the ridgeback. vessel. CPUE is measured as pounds harvested per

19 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

TABLE 8 1990. The objective of these new measures will be Sea Urchin Landings (1,000s of Pounds) to further reduce harvesting pressure, especially in Northern Southern northern California. Resource surveys and fishery Year California California Total monitoring programs will continue and will be ~__~ ~~ - ~__ 1971 0 <1 <1 increasingly important for evaluating manage- 1972 <1 76 76 ment changes. 1973 18 3,594 3,612 1974 51 7,056 7,107 1975 3 7,323 7,326 RECREATIONAL FISHERY 1976 95 11,012 11,107 1977 386 16,208 16,594 Catches from the California commercial passen- 1978 34 14,394 14,428 ger (CPFV, or partyboat) fleet can gen- 1979 237 20,307 20,544 erally be considered indicative of nearshore and 1980 103 21,196 21,299 1981 194 24,720 24,914 offshore sport angler success. The CPFV fleet can 1982 92 19,347 19,439 locate and catch any species available within the fish- 1983 61 17,207 17,268 ing area. Catches can vary widely for latitudinally 1984 59 14,920 14,979 1985 1,921 18,074 19,995 migratory species, such as barracuda (Sphyraena ar- 1986 10,174 23,957 34,131 genteu) and yellowtail (Seviola lalandei), and for 1987 23,600 22,500 46,100 highly migratory transoceanic species like albacore. 1988 30,525 21,463 51,988 1989* 26.745 24,168 50,913 Catches of resident species in nearshore areas may

~~ ~~ *Preliminary also show fluctuations associated with warmer oceanic regimes. Partyboat landings for 1989 - 4.4 million fish - diving hour. The northern California average was were slightly higher than in 1988. Rockfish main- 570 pounds per hour in 1989, compared to 505 tained its first-rank position, with 2.1 million pounds per hour in 1988. In southern California the fish caught; this is about a 15% increase over 1988 1989 average CPUE was 323 pounds per hour, rang- (table 9). ing from 166 at the Palos Verdes Peninsula to 516 at Sand bass (Pavalabvax nebtili$er) landings again ex- San Nicolas Island; in 1988, the average was 286 ceeded kelp bass (Pavalubrax clathratus) landings, pounds per hour and ranged from 160 to 393. which increased 16% over 1988. Sculpin (Scorpaena Logbook data show that the majority of harvest- gtittata) maintained its sixth-place ranking: 25% ing effort in northern California occurred between more fish were caught than in 1988. The barracuda Fort Bragg and Gualala (figure l), but increased ef- fort also took place in the Westport area to the north and at the Farallon Islands to the south. Over 60% TABLE 9 of the effort in southern California was expended at 1989 Commercial Passenger Fishing Vessel Catch the Channel Islands, with 37% at the four northern islands and 17% at three of the southern islands. The Specieslspecies Thousands of fish Rank San Diego coastal area was the highest mainland group Rockfish 2,135 1 zone, receiving 15% of the effort. Sandbass 415 2 Size distributions of sea urchins landed in north- Kelp bass 373 3 ern California have changed slightly, with a mean Pacific mackerel 350 4 Bonito 322 5 size of 103 mm (108 mm in 1988). Only 3.6% ofthe Sculpin 161 6 samples were smaller than the 76-mm minimum Barracuda 133 7 size, which was adopted in March 1989. This new Salmon 110 8 Lingcod 75 9 size regulation appears to have affected harvesting Halfmoon 67 10 practices in southern California. The mean size of Yellowtail 61 11 sampled sea urchins was 94 mm (91 mm in 1988), Ocean whitefish 44 12 Albacore 29 13 and the overall percentage of sea urchins below the Flatfish (mix.) 28 14 minimum size dropped from 17% in 1988 to 10% Sheephead -_77 15 in 1989. In coastal areas such as Santa Barbara and Skipjack tuna 20 16 Yellowfin tuna 17 17 the Palos Verdes Peninsula, percentages of undersize White croaker 15 18 sea urchins decreased from as high as 38% in 1988 California halibut 9 19 to as low as 10% in 1989. Bluefin tuna 6 20 Others 61 - The sea urchin fishery is likely to come under ~~ . increasingly restrictive management measures in Total 4.453

20 FISHERIES REVIEW: 1989 CalCOFl Rep., Vol. 31,1990

catch was still well above the lower catches in the 15 Contributors: years before 1987. Salmon (Oncorhynchtis sp.) also Kristine Barsky, ridgeback and spot prawns had a good year. The lingcod (Ophiodon elongatus) Patrick Collie5 Pacijic Ocean shrimp take increased 19%. Albacore finally made the top Cedric Cooney, northern anchovy 20, although mostly small fish in the eight-pound Gary Galovich, Pacific sardine range were caught. The highly desirable California Frank Hen ry, groti ndjsh halibut retained nineteenth place with only 9,000 Mary Larson, albacore fish, a 25% decrease from 1988. Striped bass (Roccns Robert Leos, market squid saxatilis) had a poor year: a little over 2,000 fish were Malcolm Oliphant, recreationaljishery caught, as opposed to over 10,000 in 1988. David Parke,; sea iirrhin Jerome Spratt, Pacific herring John Sunada, swordjish and shark Phillip Swartzel I, Cal ijornia spiny lobster Patricia Velez, Calijornia halibtit Ronald Warner, Dungeness crab Patricia Wolf; Pacific mackerel

Compiled by Tervi Dickersoti

21 PUBLICATIONS CalCOFl Rep., Vol. 31,1990

P UB 1 I CAT1 0 N S January 1 through December 31,1989

Balch, W., R. E. Eppley, M. R. Abbott, and F. M. H. Reid. Bias in 1986-87 fishing year. Final Rep. Coop. Agreement NA-86-ABH- satellite-derived pigment measurements due to coccolithophores and 00018. NOAAiNMFS SWR, January 1989, 10 pp. dinoflagellates. J. Plankton Res. 11(3):575-581. Hanan, D. A,, L. M. Jones, and R. B. Read. California interac- Baumgartner, T. R., J. Michaelsen, L. Thompson, G. Shen, A. Soutar, tion and depredation rates with the commercial passenger fishing and R. Casey. The recording of interannual climatic coral bands, ice vessel fleet near San Diego. Calif. Coop. Oceanic Fish. Invest. Rep. cores and varved sediments. In Aspects of climatic change over the 30: 122-1 26. eastern Pacific and western Americas, D. Peterson, ed. Amer. Geo- Hanan, D. A , J. P. Scholl, and S. L. Diamond. Harbor seal, Phoca phys. Union, Geophys. Monogr. 55:l-14. virditla vichardsi, census in California, May-June 1988. January 1989. Beers, J. R., and D. C. Brownlee. Laboratory observations on the re- SWFC Admin. Rep. #LJ-89-13, 53 pp. production and feeding of a marine pelagic strombidiid ciliate. Un- Hunter, J. R., B. J. Macewicz, and C. A. Kimbrell. Fecundity and other versity of California, Institute of Marine Resources, Reference aspects of the reproduction of sablefish, Anoplopomajimbvia, in central 88-4.25 pp. California waters. Calif. Coop. Oceanic Fish. Invest. Rep. 30:61-72. Boveng, l? L., A. A. Hohn, D. P. DeMaster, A. E. Dizon, D. A. Hanan, Jackson, G. A,, F. Azam, A. F. Carlucci, R. W. Eppley, B. Finney, D. S. andJ. l? Scholl. Tracking harbor seals in Southern California. In Proc. Gorsline, B. Hickey, C. Huh, R. A. Jahnke, I. R. Kaplan, M. L. 1989 North Amer. Argos users conf. and exhibit. Service Argos, Inc., Landry, L. F. Small, M. I. Venkatesan, P. M. Williams, and K. M. 1801 McCormick Dr., Suite 10, Landover, MD., 20785, pp. 205-217. Wong. Elemental cycling and fluxes off the coast ofsouthern Califor- Butler, J. L. Growth during the larval andjuvenile stages ofthe northern nia. EOS 70:146-149, 154-155. anchovy, Engradis movdax, in the California Current during 1980- Jacobson, L. D., and N. C. H. Lo. Spawning biomass of the northern 84. Fish. Bull. 87(3):645-652. anchovy in 1989. SWFC Admin. Rep. #LJ-89-17. Butler, J. L., C. A. Kimbrell, W. C. Flerx, and R. D. Methot. The Jacobson, L. D., and C. J. Thomson..Evaluation of options for manag- 1987-88 demersal fish surveys off central California (34"30'N). U. S. ing northern anchovy- a simulation model. SWFC Admin. Rep. Dept. Commer., NOAA Tech. Memo., NMFS, SWFC, No. 133, #LJ-89-26. 44 PP. Kalvass, P. Characteristics of red sea urchin, Stvongylocentvotusfrancis- Chereskin, T. K., E. Firing, and J. A. Gast. On identifying and screen- mitis, populations in the vicinity of Fort Bragg, California. Calif. ing filter skew and noise bias in acoustic Doppler current profiler Dept. Fish. Game, Mar. Res. Admin. Rep. 89-1. measurements. J. Atmos. Oceanic Tech. 6:1040-1054. Kimor, B., and F. M. H. Reid. Large dinoflagellates of the orders Gym- Chereskin, T. K., M. D. Levine, A. J. Harding, and L. A. Regier. tioditiial~s,Noctilucales and Pyrocystales from the west coast of North Observations of near-inertial waves in acoustic Doppler current pro- America. IMR Ref. 89-1. filer measurements made during MILDEX. J. Geophys. Res. Lea, R. N., J. M. Duffy, and K. E. Wilson. The Cortez angelfish, 94~8135-8145. Pomacarithrrs zotiipectus, recorded from southern California. Calif. Chereskin, T. K., €? F? Niiler, and P. M. Poulain. A numerical study of Dept. Fish Game 75(1):45-47. the effect of upper ocean shear on flexible drogued drifters. J. Atmos. Lea, R. N., K. A. Karpov, and L. E Quirollo. Record of roughscale Oceanic Tech. 6243-253. sole, CIidodevma aspervimnm, from northern California with a note on Craven, D. B., and A. F. Carlucci. Spring and fall microheterotrophic the Pacific lined sole, Aclzivus mezatlarins. Calif. Dept. Fish Game utilization of dissolved free amino acids in a California borderland 75(4):239-241. basin. Mar. Ecol. Prog. Ser. 51229-235. Leong, R. J. H. Sperm concentrations and egg fertilization rates during Dayton, P. K. Interdecadal variation in an antarctic sponge and its pred- spawning of captive anchovy, Eqvacilir movdax. Calif Coop. Oceanic ators from oceanographic climate shifts. Science 245:1484-1486. Fish. Invest. Rep. 30:136-139. Dayton, P. K., J. H. Carleton, A. G. Mackley, and l? W. Sammarco. Lo, N. C. H. Evaluation of the port sampling of the wet fish fishery. Patterns of settlement, survival and growth ofoysters across the Great SWFC Admin. Rep. #LJ-89-10. Barrier Reef. Mar. Ecol. Prog. Ser. 54:75-90. Lo, N. C. H., J. R. Hunter, and R. P. Hewitt. Precision and bias of Dayton, P. K., R. J. Seymour, P. E. Parnell, and M. J. Tegner. Unusual estimates oflarval mortality. Fish. Bull. 87(3):399-416. marine erosion in San Diego County from a single storm. Estuarine Lo, N. C. H., and R. D. Methot. Spawning biomass of the northern Coastal ShelfSci. 29:151-160. anchovy in 1988. Calif. Coop. Oceanic Fish. Invest. Rep. 30:18-31. Dayton, P. K., and M. J. Tegner. Bottoms below troubled waters: McGowan, J. A. Pelagic ecology and Pacific climate. In Aspects benthic impacts of the 1982-1984 El Niiio in the temperate zone. In of climate variability in the Pacific and the western Americas, Ecological consequences of the 1982-1983 El Niiio to marine life, D. H. Peterson, ed. Amer. Geophys. Union, Geophys. Monogr. P. W. Glynn, ed. Elsevier Oceanography Series. 55:141-150. Dillon, T. M., J. N. Moum, T. K. Chereskin, and D. R. Caldwell. On Methot, R. D. Synthetic estimates of historical abundance and mortal- the zonal momentum balance at the equator. J. Phys. Oceanogr. ity for northern anchovy. In Mathematical analysis of fish stock dy- 19:561-570. namics, E. F. Edwards and B. A. Megrey, eds. Am. Fish. Soc. Symp. Ebert, E., and J. L. Houk. Abalone cultivation methods used at the Ser. No. 6, pp. 66-82. California Department of Fish and Game's Marine Resources Labo- Mullin, M. M., E. R. Brooks, and E. F. Stewart. Nearshore surface- ratory. In Handbook of culture of abalone and other marine gastro- dwelling zooplanktonic assemblages off southern California during pods, K. 0. Hahn, ed. CRC Press, pp. 239-254. the anomalous winters of 1983 and 1984. Cont. ShelfRes. 9:19-36. Forney, K. A,, D. A. Hanan, and J. Barlow. Detecting trends in harbor Mullin, M. M., and A. Conversi. Biomasses ofeuphausiids and smaller porpoise abundance using aerial surveys: a preliminary report based zooplankton in the California Current - geographic and interannual on three years. August 1989. SWFC Admin. Rep. #LJ-89-20. comparisons relative to the Pacific whiting, Merlticciuspvodrrctzis, fish- Graham, J. B., W. R. Lowell, N. C. Lai, and R. M. Laurs. O2tension, ery. Fish. Bull. 87:633-644. swimming-velocity, and thermal effects on the metabolic rate of the Niiler, I? P., M. R. G. Poulain, and L. R. Haury. Synoptic three-dimen- Pacific albacore, Thurzriur alalurzga. Exp. Biol. 48:89-94. sional circulation in an onshore flowing filament of the California Hanan, D. A. Harbor porpoise, Pliocoem plzocoem, aerial surveys, Sep- Current. Deep-sea Res. 36:385-405. tember 1988 to November 1988. Final Rep. Coop. Agreement NA- Oda, K. T. The Pacific herring, Clupeu harmprpallasi, open pound roe- 86-ABH-00018. NOAA/NMFS SWR, March 1989, 30 pp. on-kelp experimental fishery in San Francisco Bay, 1988 to 1989. Hanan, D. A,, and S. L. Diamond. Estimates of sea lion, harbor seal, Calif. Dept. Fish Game, Mar. Res. Admin. Rep. 89-7. and harbor porpoise mortalities in California set net fisheries for the Ohman, M. D., J. M. Bradford, and J. B. Jillett. Seasonal growth and

22 PUBLICATIONS ColCOFl Rep., Vol. 31,1990

lipid storage of the circumglobal, subantarctic , Neocalaruis national Symposium on Operational Fisheries Oceanography toizsus (Brady).Deep-sea Res. 36:1309-1326. (ISOFO), St. John's, Newfoundland, Canada. 23-27 October 1989. Ohman, M. D., and J. R. Wilkinson. Comparative standing stocks of Small, L. W., M. R. Landry, R. W. Eppley, E Azam, and A. E Carlucci. mesozooplankton and macrozooplankton in the southern sector of Role of plankton in the carbon budget of Santa Monica Basin. Mar. the California Current system. Fish. Bull. 87(4):967-976. Ecol. Prog. Ser. 53:57-74. Owen, R. W. Microscale and finescale variations of small plankton in Smith, E., M. D. Ohman, and L. E. Eber. Analysis ofthe patterns of coastal and pelagic environments. J. Mar. Res. 47:197-240. distribution of zooplankton aggregations from an acoustic Doppler Owen, R. W., N. C. H. Lo, J. L. Butler, G. H. Theilacker, A. Alvarifio, current profiler. Calif. Coop. Oceanic Fish. Invest. Rep. 30:88-103. J. R. Hunter, and Y Watanabe. Spawning and survival patterns of Spratt, J. D. Biological characteristics of the gillnet catch from the larval northern anchovy, Erzgvaulis movdax, in contrasting environ- 1988-89 Pacific herring, Clupea hareips yallasi, roe fishery in Cali- ments - a site-intensive study. Fish. Bull. 87(3):673-688. fornia. Calif. Dept. Fish Game, Mar. Res. Admin. Rep. 89-5. Parks, W. W., S. E Herrick, Jr., and P. J. Donley. U.S. trade in tuna for . Biomass estimates of Pacific herring, Clupea hareizglrspallasi, in canning, 1987. Southwest Region Admin. Rep. #SWR-89-01. California from the 1988-89 spawning-ground surveys. Calif. Dept. Parrish, R. H., N. W. Bartoo, S. F. Herrick, Jr., P. M. Kleiber, R. M. Fish Game, Mar. Res. Admin. Rep. 89-6. Laurs, and J. A. Wetherall. Albacore management information doc- -. The distribution and density of eelgrass, Zosteva mavirza, in ument. U.S. Dept. Commer., NOAATech. Memo., NMFS, SWFC, Tomales Bay, California. Calif. Dept. Fish Game 75(4):204-212. No. 126, 56 pp. Squires, D., S. F. Herrick, Jr., and J. Hastie. Integration ofJapanese and Porter, R. G., B. Culver, I. H. Hreha, and M. L. Larson. Pacific coast United States sablefish markets. Fish. Bull. 87(2):341-351. fishery review reports, albacore fishery in 1988-Appendix 2. Irz 41st Stevens, E. G., W. Watson, and H. G. Maser. Development and distri- Ann. Rep. Pac. Mar. Fish. Comm., R. G. Porter, ed. Pp. 9-10. bution of larvae and pelagicjuveniles of three kyphosid fishes (Girdla Reid, E M. H., and E. Stewart. Nearshore microplanktonic assem- rzipicarir, Medidrtrza cal$iovrziensis, and Hevmosilla azurea) off California blages off southern California in February 1983 during the El Nifio and Baja California. Fish. Bull. 87(3):745-768. event. Cont. ShelfRes. 9(1):37-50. Tegner, M. J. The California abalone fishery: production, ecological Reid, J. L. Circumpolar water in the North Atlantic. EOS 70(43):1132 interactions and prospects for the future. 1rz Scientific approaches to (abstract). management of invertebrate stocks, J. F. Caddy, ed. New York: John . On the total geostrophic circulation of the South Atlantic Wiley and sons. Pp. 401-420. Ocean: flow patterns, tracers and transports. Prog. Oceanogr. Tegner, M. J.. P. A. Breen, and C. E. Lennert. Population dynamics of 23(3):149-244. red abalones (Haliotis vufesmis) in southern California and manage- Reilly, P N., K. T. Oda, and E E. Wendell. Pacific herring, Clirpea ment of the red and pink (H. covnrgata) abalone fisheries. Fish. Bull. harerzgur pallasi, studies in San Francisco and Tomales bays, April 87(2):313-339. 1988 to March 1989. Calif. Dept. Fish Game, Mar. Res. Admin. Tegner, M. J., and R. A. Butler. Abalone seeding. In Handbook of Rep. 89-4. culture of abalone and other gastropods, K. Hahn ed. Boca Raton, Riddle, M. J., D. M. Alongi, P. K. Dayton, J. A. Hansen, and D. W. Florida: CRC Press. Pp. 157-182. Klumpp. Detrital pathways in a coral reef lagoon. Mar. BIOI. Theilacker, G. H., and Y Watatiabe. Midgut cell height defines nutri- 104:109-118. tional status of laboratory raised larval northern anchovy, Eriguaulis Roemmich, D. Mean transport of mass, heat, salt and nutrients in movdax. Fish. Bull. 87(3):457-469. southern California coastal waters: implications for primary produc- Thomson, C., C. Cooney, and J. Morgan. Status of the California tion and nutrient cycling. Deep-sea Res. 36(9):1359-1378. coastal pelagic fisheries in 1988. SWFC Admin. Rep. #lJ-89-14, Roemmich, D., and T. McCallister. Large scale circulation of the North 25 pp. Pacific Ocean. Prog. Oceanogr. 22:171-204. Venrick, E. L. Behind a front: an overview of the Ensenada Front Proj- Seymonr, R. J., M. J. Tegner, I? K. Dayton, and I? E. Parnell. Storm ect. EOS 71(2). wave induced mortality of giant kelp, Macvocystispyrijera, in southern Wahl, D. D., and J. J. Simpson. Velocity estimates from space: sensitiv- California. Estuarine Coastal Shelf Sci. 28277-292. ity to computational methods and near-surface physical processes. Simpson, J. J. An integrated study of offshore mesoscale eddy dynamics IEEE/Oceans '89 Conference Proceedings, 18-21 September 1989, in the California Current system. IEEE/Oceans '89 Conference Pro- Seattle, Wash. ceedings, 18-21 September 1989, Seattle, Wash. Watson, W., and R. I. Davis, Jr. Larval fish diets in shallow coastal . Remote sensing in fisheries: a tool for better management in waters off Sail Onofrc, California. Fish. Bull. 87(3):569-591. the utilization of a renewable resource. Symposium abstracts, Inter-

23

SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT ColCOFl Rep., Vol. 31,1990

Part II FORTIETH ANNIVERSARY SYMPOSIUM OF THE CALCOFI CONFERENCE La Jolla, California October 27,1989 OCEAN OUTLOOK: GLOBAL CHANGE AND THE MARINE ENVIRONMENT

The fortieth anniversary symposium was held on who is the chairman of the Board of Regents of the the final day of the 1989 CalCOFI Conference. The University of California. We are clearly delighted morning session featured an award to Roger Revelle that he has taken the time to come down and listen and talks by Lieutenant Governor Leo McCarthy to what we’re doing here. I’d also like to welcome and Under Secretary of Commerce for Oceans and Claire Dedrick, executive director of the State Lands Atmosphere John Knauss. The afternoon session Commission; Mike McCollum, chief deputy direc- was devoted to a panel discussion of the question tor of the California Department of Fish and Game; “What does society need from ocean scientists in and Mario Martinez, director of our sister university preparing for global change?” Panelists were seven system in Mexico, CICESE. We have strong and distinguished representatives of scientific, govern- ongoing relations across the border; we try very mental, and environmental agencies. hard to maintain them and keep up our end of the Proceedings for both sessions were recorded, bargain. I think it’s extremely important that we do transcribed, edited for clarity, and reviewed by the this. I’d also like to welcome Howard Ness, who is speakers. We trust that the transcribed talks will be the U. S. regional fisheries attach6 for the American as informative and enjoyable to read as they were to embassy in Mexico City; Craig Denisoff, who is hear. representing California Senator Barry Keene; Alicia The CalCOFI Committee Wen bourne, rep re sent ing Assemblyman Gerald Felando; and representatives of the office of City Edward A. Frieman: I’d like to welcome all of you Councilman Bruce Henderson. There are probably on behalf of the faculty, staff, and students of the others here whom I should have recognized and Scripps Institution of Oceanography of the Univer- didn’t; my apologies. sity of California, San Diego. I’d like to express our Of course a very special welcome to - I guess I’d appreciation for CalCOFI’s selecting our campus for have to call him a Renaissance man - Roger Revelle. the location of this fortieth anniversary celebration. This conference comes at a difficult time in our It gives an opportunity for many of us to hear and history, when humankind is being stressed by severe participate in what you are doing and, of course, to environmental change. And clearly the combined allow members of the San Diego community to join efforts ofscience and society will be required to meet in. this challenge. It seems to me that there’s a growing I’d also like to give special greetings to a few peo- appreciation of the strong linkages that exist be- ple. I really appreciate the presence of the Honorable tween environmental policy, energy policymaking, Leo McCarthy, lieutenant governor of California, and economic policymaking throughout many sec- and of the Honorable John Knauss, who is the under tors of society. There’s a growing realization in the secretary of commerce for oceans and atmosphere, international arena that action is needed. We see and, as it’s been pointed out many times, one of us - Maggie Thatcher and President Mitterand propos- a Scripps graduate. John still has a home here, so I ing major conferences and international action in think we can claim some special attention on his this regard. And there’s a slower-growing realiza- time. tion that a strong science base is needed, much more There are many others I would like to welcome. I so than in the past, to support this kind of policy- would like to give a special welcome to Roy Brophy, making. So it seems very fitting that CalCOFI

25 SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990

sponsor a symposium on global change, bringing CalCOFI, as you know, was a response to a de- together scientists, government leaders, and those cline in the catch of the sardine fishery, which was at with environmental concerns to focus attention on that time the largest single fishery in the world. I’m these matters of science and public policy. told the sardine fishery is back, at least to the degree It’s also noteworthy that part of our understand- that fish have been canned again in 1988. Clearly the ing of the impact that the oceans have on climate program would like to take credit for that, and came from CalCOFI, whose primary goal is under- though you are great, I think you don’t yet walk on standing the marine life of the California Current. water. But you did provide the understanding that CalCOFI has led in many areas of research on air- led to much of this. sea interaction, the dynamics of ocean currents, the An issue then is the role of CalCOFI in the future, El Nifio phenomenon, and other insights vital to our and I look to CalCOFI for major leadership. I think understanding of both short-lived and long-lived, there is a growing understanding throughout the short-range and long-range weather predictions. oceanographic community of a new kind of unifi- I’m told that there was a CalCOFI meeting in the cation of a number of disciplines - chemical, bio- late 1950s that brought together some of the nation’s logical, physical, marine biological - to attack the leaders in ocean and atmospheric science, including global problems we must deal with in the future. many from Scripps, to discuss the role of the oceans We can no longer maintain the stance in the scientific in determining climate. literature, in our research, that these areas are dis- It’s my contention that those in the political arena, tinct and separate. For example, there is talk in the worrying about the issues of global warming, tend National Science Foundation of a new initiative in to think of it purely as the province of the atmos- global ecosystems dynamics, called GLOBEC. I pheric sciences. The major role of the oceans, which would hope that the Marine Life Research Group perhaps will be discussed today in later sessions, is and CalCOFI will play a major role in that new en- often disregarded, poorly understood, swept aside. deavor when and if it gets started. It is not part of the national agenda in this debate, Lastly, I would like to commend the CalCOFI and I think one of the outcomes of meetings like this Committee for their really tireless efforts over the will be to help us focus on this issue, which is a very past year to provide this forum. There are many serious issue for the future. people on the planning committee - Izadore Bar- An outgrowth of the work of CalCOFI was the rett, who is director ofthe Southwest Fisheries Cen- establishment of the Scripps Center for Climate Re- ter of NOAA’s National Marine Fisheries Service; search. Later on, the nation’s first experimental Richard Klingbeil of the California Department of long-range forecasting center was established at Fish and Game; the current CalCOFI coordinator, Scripps in the 1970s. CalCOFI has assembled an Patricia Wolf, from the California Department of enormous data base on ocean temperature, salinity, Fish and Game; and last, but certainly not least, and circulation patterns, along with comprehensive Mike Mullin, director of our Scripps Marine Life marine life collections from larvae to adult fishes and Research Group; and his assistant George Heming- including marine mammals and birds. This is all way. I know, from Mike’s first talking to me about invaluable to our research in global change. We must this, many months ago, that he has been a superb have data of high quality and continuity for project- leader, tireless in his efforts on your behalf. I wish to ing changes in climate and species. thank you all. It has been noted that CalCOFI began in the Finally, on behalf of the institution, I would like 1940s. It is really amazing in the world of science for to welcome you to what promises to be an exciting such a cooperative program, which involves federal day of talk and debate as we explore the new roles of and state governments, universities, and support science and public policy in meeting this challenge. from the private sector, to be able to function for four decades. The relaxation time for political phe- Michael M. Mullin: I want to add my welcome to nomena tends to be four years, yet we have func- that of Professor Frieman, and I stress that this is a tioned and stayed together over many, many triple welcome, because, as you know, one of the political decades. I cannot think of another major strengths of the California Cooperative Oceanic scientific program that has this characteristic. From Fisheries Investigations is the enduring partnership the scientific point of view, one might say it provides between the Marine Life Research Group here at evidence that it can be done. I think it’s important Scripps; the Southwest Fisheries Center of the Na- for policymakers at high levels to be aware ofthat. tional Marine Fisheries Service, National Oceanic SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990

and Atmospheric Administration; and the Califor- was sampled each month. The second slide (Hewitt’s nia Department of Fish and Game. Therefore, wel- figure 6) represents the years from 1961 through come, welcome, and welcome. 1965. Coverage was reduced to approximately This symposium takes place on the third day of quarterly, but was still spatially extensive. The third the annual scientific meeting of this consortium; to- slide (Hewitt’s figure 7) is for 1966 through 1978, gether with other interested scientists, we’ve been when extensive coverage was reduced to every third sharing our developing, hard-won knowledge and year. The final slide (Hewitt’s figure 9) represents speculations. The symposium today is really about 1979 through 1987. The weakness of the triennial the future - about the problems that may arise for system was revealed in 1983, when a major El NiAo society from environmental change, and the roles of had the impudence to occur in a non-CalCOFI year. marine scientists in addressing these problems. Resources were begged, borrowed, and stolen to set We will hear from distinguished and diverse sci- up a single line off Del Mar, California, and in 1984 entists and policymakers as they discuss and debate the sampling plan was changed to quarterly cover- these issues from several perspectives, and we are age of the segment from San Diego to San Luis very grateful for their participation. I particularly Obispo. And that is the pattern that continues today. want to thank our participants from the Bay Area There are sound scientific reasons for reducing the and Sacramento - Lieutenant Governor McCarthy, area of coverage, as well as financial pressures, but Assemblyman Sher, and Professor Scheiber - who I’ll spare you the arguments. are with us in spite of the recent earthquake and its I don’t want to leave you with the impression, major disruption of normal life. though, that all of the seagoing science has been Yet the symposium also honors the past - forty confined to the California Current. The basic meth- years of a collaborative effort to develop an under- ods have been used by CalCOFI scientists and others standing of the sea off California and its living re- to map most of the North Pacific. Furthermore, not sources. Put quite simply, we believe that CalCOFI shown on these maps are extensive and intensive exemplifies the three bases for grappling with future cruises within the current, specifically designed to environmental problems. These are the fund of fac- assess the abundance of eggs and larvae of commer- tual knowledge that has been accumulated; the con- cial species of fish. ceptual and technical tools that have been developed The second visual image is a videotape that Chuck (and which, incidentally, have been adopted in Colgan and Bill Call of Scripps prepared for this many parts of the world); and the model of cooper- anniversary celebration, starring one of our own re- ation and mutual support between organizations searchers. We plan to distribute this widely as a whose internal politics are often quite divergent. We source of information about CalCOFI and about believe, to paraphrase Santayana’s oft-quoted re- how large-scale issues in the marine environment mark about history, that those who do not under- have been tackled. (videotape) stand the environmental change of the past are As you can see, we are very proud of the cooper- condemned to misinterpret what is happening to ative aspect of this program, and although it’s cer- them in the present. tainly evident to those of us in it, I should point out As proud as we are of the record of this pro- to newcomers that there has also been a very long- gram-and we all are proud of it-we organized standing and fruitful cooperation with marine re- this symposium for the future, so I would like to searchers and fisheries biologists in Mexico. They leave you with some visual images representing the are regular participants in the CalCOFI conference, CalCOFI program up to this point. and we welcome their continuing involvement, be- First, to demonstrate that the program has been cause the problems that we face simply don’t rec- solidly based in seagoing, though by no means lim- ognize national borders. ited to that, Roger Hewitt’ put together maps of the Finally, to remind you again of the reason for this coverage of the California Current by the CalCOFI symposium in words from outside CalCOFI, I cruises since 1949. I show these to you for an overall quote from Our Changing Planet, An Executive Sum- visual impact, not for details. The first slide (Hew- mary ofthe U.S. Global Change Research Program: itt’s figure 5) shows the coverage from 1949 through 1960, during which much of the California Current Although human activities may have the potential to alter the Earth system, it is clear that variations occur

‘Historical review of the oceanographic approach to fishery research, Calif. naturally over a wide range. For many of these changes, Coop. Oceanic Fish. Invest. Rep. 29:27-41. current knowledge is insufficient to reliably predict the

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likely debate, rate, or timing of these changes. To under- vessels Horizon and Crest and the Cal Fish and Game stand and ultimately predict the impact of both natural vessel N.B.Scojeld were able to go to sea. Much of processes and human activities on these changes, it is the work on the first few cruises was done by stu- necessary to improve our understanding of the underly- dents and at least one professor, Norris Rakestraw ing physical, geological, chemical, biological, and social (he was on cruise 1 when I was). processes that control the earth’s environment. . . . An Well, Roger supported this Marine Life Research effective and well-coordinated national and international research program will be required to dramatically im- Program of the Scripps Institution, a component of prove our knowledge of these complex earth processes - CalCOFI and its sardine study, arguing with his to provide the basis to discriminate between natural and characteristic vigor for a type of environmental ap- man-influenced changes and ultimately to predict global proach now called fisheries oceanography. He pur- change. sued his aims with so much vigor that some of the non-Scripps scientists of the time complained that, So much for background. My distinguished pred- lacking the mandates that the federal and state agen- ecessor as director of the Marine Life Research cies had, Roger was sometimes insensitive to their Group was Professor Joseph Reid, who led the uni- concerns and interagency rivalries. versity’s part of the CalCOFI program for many Roger became director in 1950, and he was, with years. Joe is, as most of you know, a physical ocean- Carl Hubbs, the consistent Scripps presence on the ographer, but he has published on chemistry and Marine Research Committee, which managed the biology as well, and represents an integrative, large- CalCOFI program. He remained actively involved scale view of the ocean. I have asked Joe to make a in that committee until 1959. Without losing sight special presentation to Roger Revelle, one of the of the concern for the economically important, but great men in CalCOFI’s history and in American failing, sardine industry, Roger was a vocal and elo- environmental science. quent proponent of the central idea of modern ecol- ogy: that any particular species is part of a physical Joseph Reid: Hello, Roger. At the time the Cal- and biological environment and must be studied COFI program was first conceived, Roger wasn’t from the broad perspective by experts in a variety of here. That was in the middle forties; I believe he was fields. He combined broad vision with great energy off as a sailor in Washington at that time. But a little and persuasive powers. He was certainly one of the later he did manage to get two ships - stealing ships first marine scientists to become interested in the from the navy in 1944 would have been awkward, possibility that adding carbon dioxide to the atmo- but in 1947 or 1948 it was a little easier; there was a sphere from the burning of fossil fuels might cause surplus. And he did this so quickly that the univer- global warming - the greenhouse effect. sity president at that time, Gordon Sproul, com- This is the fortieth anniversary of CalCOFI, but I plained to Harald Sverdrup that he’d only learned was reminded by Saul Alvarez-Borrego yesterday by memorandum after the event that he was now that it’s also the fiftieth anniversary of your trip, responsible for a fleet. Roger, on the E. W Scripps to the Gulf of California Well, getting ships out of the navy at that time in 1939. And that merits some mention too. was the easiest part of the job. When Roger came to I have heard, and it’s only a rumor, that while the Scripps as associate director in 1948, he had to use ship was down there and you were engaged on your them. One of his many responsibilities was to see work on the sediments in some of the basins, the that these ships were equipped and manned with captain had to leave, and you brought the E.W people who knew how to carry out the work at sea. Scripps back to San Diego. I don’t know whether This was much the hardest part of the job. this is true or not, but it’s rather frightening to think In the earlier part of the Scripps career, the faculty of the scientific leader of a cruise all of a sudden had done much of their own work at sea. Because becoming the navigator. Those of you who have most of the cruises were fairly short, this had been to sea know what sort of scientists we’ve got. worked out well. But monthly cruises of three ships All are very competent in their own particular fields, were a different order, of course. And they would which may be deep, but not necessarily very wide. need more trained people. Starting, I believe, with a I’m afraid that there are some of us who, given the core of one experienced marine technician in 1948, problem ofnavigating a ship, once we’ve gotten off- and enough gear for one ship, Roger somehow man- shore far enough not to see North America, couldn’t aged to find enough instruments and trained people find it again. . . . And there are some of us who can for three ships. So that in March of 1949 the Scripps do anything. I don’t know whether Roger really

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brought that ship back by himself, but in his case, I son. And although Joe gives me the credit for don’t think there’s any doubt that if he’d put his thinking about the problem from an ecological hand to it, he could have done that, as well. point of view, Harald, in spite of the fact that he was Today, as a symbol of our recognition of what he’s primarily a geophysicist, also had ecological ideas, done for this program, we present Roger with a very much so. scale model of the Scripps vessel Ellen B. Scripps, The other man who pushed CalCOFI very hard which has been used for research in the California and had many original insights was John Isaacs, who waters. This model was made by George Snyder, was director of the Marine Life Research Group for who has cared for the plankton collections at Scripps several years. I think that was perhaps the happiest for many years. He has received, cataloged, and pre- time of his life. He was a biologist manquk. He was served plankton samples collected by Scripps vessels trained as an engineer and a physicist, but all his life from all the world’s oceans. And he certainly knows he wanted to be a biologist. And to a considerable what goes on the fantail of a Scripps vessel. This is extent he wus a biologist. One of the most interest- not the largest or the newest of the Scripps vessels. ing things he did was to start a program of coring in It’s a workhorse for much of the research Scripps has the Santa Cruz Basin, south and west of Santa Bar- carried out in the inshore waters. A small token, but bara, where there are varved sediments deposited given sincerely. Thank you, Roger. under anaerobic conditions. The study of the fish remains in those sediments showed that long before Roger Revelle: Thank you very much, Joe. And there were any fishermen, any Cannery Row, any thank you all. John Steinbeck, any collection of Portuguese and This occasion brings back a flood of memories of Italian fishermen in California, the sardines fluc- the early days of this remarkable program. It is quite tuatedjust about as much over several hundred years right that we went to the Gulf of California in 1938. as they have since 1930..And there was an alternation Harald Sverdrup led that expedition. In 1939 Francis between the populations of sardines and of anchov- Shepard and Charlie Anderson and I led a geological ies long before any human activities affected the expedition on the old E. W Scvippr. On the way back, fishery. the engine broke down. To get out of the gulf and to Another man who was very much involved in this get along the coast of Baja California, we had to sail. program is still alive. He is Jack Marr, who’s sitting We sailed a very long tack to the westward- about right here in this room. He played a major role in the 800 miles, as I remember-and then sailed back early days of the program. John McGowan, who is again, a total distance of about 1600 miles, and we also here, was another one. made 20 miles good. (laughter) This was a very dif- What’s remarkable to me is that the CalCOFI pro- ficult way to get from Baja California to San Diego. gram is still being enthusiastically pursued after Fortunately we did have some professional sailors on more than forty years. People still think about it board (we were not all amateurs), and the sailors hard and work hard on it, and obtain interesting new could see that unless we got that engine fixed we’d results. It’s quite a remarkable scientific operation to never get home. So they doubled and redoubled and have gone on so long and so effectively. I’m very quadrupled their efforts, and pretty soon the engine proud to have been involved with it in the early days. was working again. So the last part of the trip was a Thank you very much. (applause) lot easier. In those days, the scientists literally were the sail- ..... ors - not the only sailors, there were also a few pros on board-but we all had to stand watches. We Mullin: It was a great pleasure for us to be able to stood six hours on and six off That’s really a very choose a gift so close to Roger’s seagoing abilities. I difficult kind of watch to stand. It’s awfully boring hope he can navigate this one home as well as he about 5 or 6 o’clock in the morning, particularly navigated the E. W Scripps. after you’ve been up since 12. Nothing ever really Next, it’s my honor and pleasure to introduce the happened, of course, but just staying awake was a Honorable Leo McCarthy, lieutenant governor of serious problem. California. His most notable environmental role is This occasion makes me think of some people on the State Lands Commission, but of course he who aren’t here, particularly Harald Sverdrup and has been connected with lands in California much John Isaacs. Harald was the man who really con- more intensively during the last two weeks, and ceived this CalCOFI program more any other per- we’re very happy that he could join us today. SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990

REMARKS BY THE HONORABLE LEO MCCARTHY, LIEUTENANT GOVERNOR OF CALIFORNIA

Director Frieman, Director Mullin, Under Sec- scientific community, the environmental commu- retary Knauss, Regent Chairman Brophy, Professor nity, a number of enlightened business leaders in the Revelle, and ladies and gentlemen. On the program state, and concerned citizens to challenge those it says that I have 45 minutes. This will not be a 6- forces and to reverse the damage they have caused in hour watch, and it won’t be a 45-minute watch. I the past and would continue to cause. once spoke for 32 minutes at a gathering, and have Of course oceans make up only a part of all of our never forgiven myself for that. So we’ll do it in a resources that are in jeopardy. Rain forests, home to somewhat shorter period of time. half the species of the world, are being eradicated. Professor Revelle is remarkable for many reasons. Acid rain threatens many of our most beautiful, I was told before we came into this auditorium that pristine areas. Chlorofluorocarbons are eating a hole just a couple of weeks ago he had his pacemaker in our atmosphere. And overdevelopment is taking checked out, and that he watched as they were doing a toll on open space, particularly on crucial wetlands it. Professor Revelle, when I was in the legislature, areas. especially as speaker, I had many incisions in my The scientists in this audience, better than anyone, heart, but they were never voluntary, and I never understand the environmental implications of these enjoyed watching them. phenomena. Your arguments are compelling for re- It struck me as I was sitting there in the front row ducing the use of chlorofluorocarbons, for banning that I was watching a man being honored, Roger the use of carcinogenic pesticides, for doing every- Revelle, who is one of the eminent scientists in the thing possible to prevent oil spills, and for recycling. world and has achieved so much. One of the things And hearing about the very real damage being from my childhood that I cling to as a happy mem- done to our environment, it is hard for me to believe ory is the one B that I got in a science course. (laugh- that we- we in a broad, public sense- don’t re- tev) We don’t have to mention what the other grades spond more vigorously. But we allow the damage to were. But that doesn’t diminish my feeling in any continue with only moderate change. Acid rain way that today’s symposium is critical, not only to wasn’t discovered yesterday. The Valdez disaster last celebrate this remarkable fortieth anniversary of March wasn’t our first exposure to the devastating CalCOFI, but in attempting to fuse the magnificent effects of a massive oil spill. And although global research that has been going on here- it’s one of the warming has gained considerable attention in the premier science programs in the world - with the last year, scientists have known about this phenom- making of public policy. enon for quite some time. Scientists are doing re- Being here in La Jolla today, I’m reminded ofJohn search and giving us much new knowledge. Steinbeck’s Cunnevy Row and of the journeys that So why the lack of progress? Because time and Doc Ricketts used to make from his biological lab again we lose vital battles in the political arena. Be- in Monterey down the coast to La Jolla, looking for cause every time scientists, and those enlightened specimens of marine life. I wonder what Doc Rick- businesspeople, and community leaders issue warn- etts would think if he made such a journey today as ings about an environmental hazard, someone inev- he passed mile after mile of vital, beautiful coastline itably clouds the issue by hiding behind specious pocked by a number of offshore oil platforms, as he arguments, usually disguised as economics, but watched a young lifeguard post another “no swim- often cloaking somewhat narrow economic self- ming’’ sign on the beach in Santa Monica, as he saw interest. The partnership between science, policy, the La Jolla tidepools he prized littered with plastic and community action must disperse those foggy six-pack rings, some lying free, some entwined arguments. around the beaks and throats of seabirds. I believe We work to require oil companies to take steps to Doc Ricketts would see what we see-that the prevent spills. They say that they can’t make those forces of human ignorance, arrogance, and greed requirements fit, because they would raise the cost have the power to turn the majesty of our environ- of oil and gasoline, thereby raising costs for thou- ment into a memory found only in fiction and sands of businesses and forcing layoffs. We hear that photographs. argument even after the Exxon I/aldez accident. It’s up to the policymakers, in concert with the We want to ban carcinogenic pesticides. The pub- SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990

lic hears that we shouldn’t because crop yields a dime from it - for cleanup - once that legislation would plummet and food prices would skyrocket. is enacted. Jt’s not intended to be punitive; it’s in- Or that pesticides really aren’t all that dangerous tended to be preventive. It will use an economic tool when properly applied. I’ve been visiting with to prevent disaster. And compared to the multibil- groups of farmers all over this state who on their lion-dollar damages experienced in Alaska by both own initiative have been reducing the use of pesti- the private and the public sector, it makes good sense cides and using alternatives - predators and para- economically as well as environmentally. sites, good bugs to kill the bad bugs. Together, the scientific and the environmental Let’s say we put together a stringent plan to cut communities must work with supportive elected of- back on chlorofluorocarbons. Before the ink on the ficials and those business leaders who are taking ini- plan was dry, chlorofluorocarbon apologists would tiatives, to try to harmonize economic growth and be contending that using substitutes would raise environmental sense. We must unify those elements prices and that thousands of people would lose their and present evidence to prove that many suffer eco- jobs. nomically because of a polluting company’s indiffer- We support mandating higher deposits on bever- ence or mismanagement. When a chemical refinery age bottles to increase recycling. Our opponents opposes tougher standards on toxic emissions be- claim store owners would have to hire extra people cause they say it will raise consumer prices and put to deal with the returns, and would have to reduce people out of work, let’s respond by talking about shelf space to create storage room for all the bottles. the medical costs to workers and nearby residents, Over and over again we lose because many peo- and about those who have to pay the medical costs. ple, especially those in my line of work, accept the And let’s talk about the economic losses from dam- opposition’s economic arguments over the scientific age done to buildings and cars and other kinds of and environmental arguments. The only exception property. Or let’s raise the specter of billions of dol- to this trend seems to be the brief periods following lars in costs and lawsuits that companies and their high-visibility disasters or scares. The Vuldex trag- stockholders will encounter if a Bhopal occurs in edy created a small window of opportunity for California. And no one can say it’s impossible, it doing something to prevent oil spills. The alar scare cannot happen. We know differently. When manu- may have made it easier to address the question of facturers resist banning chlorofluorocarbons for cost pesticides. But those windows may already be clos- reasons, or belittle the threat of global warming, ing. The best long-term solution is for us to increase let’s respond with costs of the fresh water and crop- our use of strong economic arguments to counter land we could lose. Or the $27 billion construction the weaker ones of our opponents. bill we’d get for new power plants to meet the in- For some of us, simply protecting the environ- creased demand for electricity. ment for our children while preventing the extinc- Our key will be credibility. Our numbers have to tion of our fellow creatures makes for a good enough be at least as believable as theirs. And our stories as argument. But others -and we must recognize clear. That’s where the scientific community must this - are moved more by economic considerations, help us again. I have suggested a fusion in the Uni- by the immediate obligation of rearing a family, ful- versity of California and other institutions, not only filling the economic obligations to dependents. the scientific community, but other departments as Those are important obligations. So we need to add well, that will look at the consequences to this state’s those considerations to the environmental and sci- future economy and to the nation’s future economy. entific arguments in terms a broader constituency You must help us translate your scientific findings can understand. The I/aldez spill cost Alaska billions into conclusions that will move the public and the of dollars in damages: lost , maybe politicians, and you must then speak loudly to those for a long time; lost tourism; and losses to other conclusions. No one will have more credibility. No- sectors of their economy as well. Those are tangible body can, when motivated to do so, speak more adverse circumstances that diminish the ability of clearly. many families to earn a livelihood. I understand that having to make this type of ar- The California State Lands Commission, which I gument can be somewhat frustrating to scientists. chair, has proposed legislation, which will be acted After all, isn’t it enough to know that chlorofluoro- upon in January, that would require oil companies carbons are raising the earth’s temperature? Isn’t it running 2,500 tanker trips up and down the Califor- enough to know that an oil spill will jeopardize the nia coast to maintain a $500 million oil spill preven- majesty ofthe coastline and the health of marine life? tion and cleanup fund. I hope we never have to spend Isn’t it enough to know that automobile emissions

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are polluting the air we breathe and shortening the children a hopelessly, needlessly damaged and dying lives of many of our fellow human beings living in planet. smoggy areas? It should be, but by themselves, these facts have not been enough...... We have to recognize which combination of ar- guments has power, and we have to make those ar- Mullin: Lieutenant Governor McCarthy may or guments. Combining your data, your conclusions may not have gotten a B in a science course, but he’s and credibility with economic projections and the obviously been studying since then. deep emotional chords that environmental issues can Our next speaker is, in a very real sense, one of strike will make the best armaments for policymak- our own -John Knauss, who is presently the under ers to take with us into the political arena. secretary of commerce for oceans and atmosphere Here in California, home to so many environ- and the administrator of NOAA. He received his mental treasures and so many environmental Ph.D. here at Scripps. He then became the distin- threats, you also have a few hard-core elected offi- guished dean of the School of Oceanography at the cials eager to work with you: Byron Sher, from University of Rhode Island. In addition to continu- whom you will hear this afternoon, is one of those. ing his interest in science, he’s been very active in Several representatives of state legislators among policy issues such as the law of the sea negotiations. that group are here this morning. Together, I know It’s a great pleasure to introduce John Knauss to we can work to win the kind of victories that ensure those few of you in the audience who don’t already we do not pass on to our children and our grand- know him.

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REMARKS BY THE HONORABLE JOHN KNAUSS, UNDER SECRETARY OF COMMERCE FOR OCEANS AND ATMOSPHERE

It is indeed a pleasure to come back to Scripps; tasks as a new employee in the Office of Naval Re- I always enjoy returning. It’s a particular pleasure to search in late 1949 was to answer a request from be invited here to celebrate an extraordinary event - Revelle to find navy loran sets that could be used on CalCOFI’s fortieth birthday - forty years of almost the research vessels involved in the program (at no continuous, systematic observations of the ocean. cost to Scripps, of course). It turned out not to be all These are not simple observations of tides or sea- that difficult to fill Roger’s request, in return for surface temperature at a few spots along the coast, which he bought me dinner at the Cosmos Club on but a complex set of physical, chemical, and biolog- his next trip to Washington. ical observations at many locations off the Pacific As Roger indicated, CalCOFI was probably the shore that require the skill, the care, and the dedica- original idea of Harald Sverdrup. CalCOFI was con- tion of many. There’s nothing else like this program ceived in the grand tradition of ICES - the Interna- on this scale in the United States, and with the pos- tional Council for the Exploration of the Sea, a sible exception of the North Sea, I’m not aware of multinational effort designed to explore the relation- anything like it anywhere else in the world. ship between fisheries and the ocean environment in We are also here to celebrate forty years of coop- the North Sea. erative work between the state and federal govern- I had an opportunity, while on sabbatical two ments and a major public university. I might note in years ago, to spend some time as an amateur histo- passing that I believe this alone makes CalCOFI a rian of marine policy and to look at how British remarkable and unusual program, particularly the fishery policy developed during its period of explo- fact that the University of California is still sive growth more than a century ago. In the process involved. I learned a bit about the early days of ICES, which, Universities have many wonderful attributes, but by the way, is celebrating its eighty-fifth birthday the care and feeding of long data sets is not one of this year. them. Universities tend to go where the action is. Although Great Britain had by far the biggest Long-term, systematic monitoring programs gen- fisheries of the North Sea, ICES was essentially a erally have a life span that corresponds to that of a Scandinavian idea. It was developed from the ideas professor. A professor retires, and suddenly the set of such scientists as Otto Petterson and Johan Hjort. of observations that he or she so carefully nurtured Their goal was to explain the fluctuations of such ceases to be a high priority of the department. Too resources as the herring fishery, the cod fishery, and often the collection ends, or its quality is allowed to the great bottom fisheries, mostly plaice, by study- erode. ing the life histories of the fish, their environmental I watched that almost happen here at Scripps, requirements, and relating those requirements to the when I was a young graduate student and Professor ever-changing physical and chemical environment McEwen retired. The question arose as to whether of the North Sea. And that was and is, of course, the Scripps would maintain his collection of surface goal of CalCOFI. Only in CalCOFI, the fish was temperatures and tide gage records taken from the originally the sardine, and the environment was the end of the pier. There was a time when even that California Current. simple set of observations was in danger of being ICES was a great success from the beginning, abandoned because no member of the faculty was but - and this is part of my story - not in its original prepared to take responsibility. mission. In retrospect, its greatest success in its first But CalCOFI has outlasted its original partici- few years was in establishing the standards of phys- pants, although a few like Joe Reid and Roger Re- ical oceanography. Through its short-lived Interna- velle continue to be actively involved in ocean- tional Hydrographic Bureau in , under the ography. I’m told that none of those involved in the leadership of - that remarkable ex- early years of CalCOFI are still part of the program. plorer, scientist, and statesman - it laid the ground- Even I can claim some indirect part of the action work for modern physical oceanography. It was during that first year of CalCOFI. One of my first there that Knudsen invented “standard seawater,”

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making it possible for technicians to measure salin- That was in 1956. So it was when CalCOFI started ity to five significant places in the cramped labora- some forty years ago, and so it was and has been for tories of small research vessels. It improved on the much of its existence. development of deep-sea reversing thermometers, These relationships are slowly yielding to analy- so they could be trusted to give accurate readings of sis. For example, one now can do a reasonablejob of in situ temperature to a few hundredths of a degree. predicting next year’s anchovy population from this It was through the bureau that Ekman developed his year’s temperature at the time of spawning. But, and equation of state of seawater, so if one knew the I expect that those of you who are professionals in temperature, salinity, and the depth at which the this room know it, understanding the detailed rela- water was taken, one could measure density to a few tionship of fisheries and hydrography continues to parts in a million. This in turn made it possible to be a challenge. calculate geostrophic currents. And of course there I would like to celebrate another aspect of Cal- was the Nansen bottle, which allowed not only the COFI. Just as laying the foundation ofmodern phys- capture of uncontaminated water at depth, but also ical oceanography was one of the unanticipated the ability to string a dozen or more such bottles achievements of ICES, so is the magnificent, unpar- on a wire and thus collect many samples simul- alleled forty-year time series of biological, chemical, taneously. and physical observations of the oceanographic con- All of this was developed by ICES in its first few ditions off the California coast a major achievement years, mostly before 1910, and these were the tech- of this program. It has produced a data set that finds niques of CalCOFI when it began operation after an increasing number of uses not related to its orig- World War 11; these were the techniques of physical inal purpose. oceanography up until about twenty or twenty-five There are two parts to this success story, and they years ago. However, ICES had less success in its are not unrelated. The first is the quality of the data. original goals, particularly in relating the physical It is not easy to maintain a tradition of quality con- environment to the abundance and distribution of trol in a complex observational program that grinds fisheries. on month after month, year after year, with an ever- ICES was originally conceived as a five-year ex- changing cast of technical observers. But it is a tra- periment. It is still going on. In 1909, at the end of dition that is absolutely essential in a program such its first five years, when the original budget was up as this. It is the tradition embodied in that wonderful for renewal, and many, including the fishing indus- Teutonic taskmaster, the late Hans Klein, who over- try, were questioning its success, Johan Hjort wrote: saw the data collection and processing program of “We hope, consequently, that the results now ob- CalCOFI for so many years. He accepted nothing tained will facilitate to a substantial extent further less than the most precise, most careful observations investigation in this difficult but yet so important a and the most rigorous analyses. field of inquiry. ” If I may be forgiven a personal note, it was a tra- Although written eighty years ago, this has a fa- dition and a program that I inherited when I began miliar ring to those of us, and I expect there are a my work on the equatorial waters, and one 1 had few in this room, who at one time or another have reason to recall a few years ago when I retired as had a large experiment not quite live up to our hopes dean at the University of Rhode Island and began and have had to go hat-in-hand to the National Sci- clearing out some old files. Among the artifacts, I ence Foundation or to the director and admit that we found an old CalCOFI form 4.5, one of those mar- might have been overly optimistic in our original velously complex plotting sheets designed by Hans proposal. (“But we are getting close, and you don’t that allowed one to plot not only the traditional want to cut off our funding now, do YOU?”) temperature-salinity relationship, but a number of But the problem of relating fisheries to hydrog- others as well, such as silicate versus thermosteric raphy was and continues to be an extraordinarily anomaly, and temperature versus oxygen, and all on difficult one. Listen to one of the best fishery biolo- at least two scales. gists of the last generation, Michael Graham, in his Time, as it does for events of one’s youth, has a excellent book Sea Fisheries, almost fifty years after way of eliminating the rough edges. I can only Hjort’s statement: “Future editions of this book dimly recall working over a light table on the equa- would certainly include more on the relations be- tor, in the days before either air conditioning or tween fisheries and hydrographic conditions, but at CTDs, sweat dripping over those damn forms the present time they are imperfectly understood.” (Hans had them printed on high-quality paper so

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that they would not disintegrate when sweated and the samples are to be trusted, and they can be upon), trying to manipulate a French curve between used with confidence. observational points. I can still remember the Hans This afternoon there will be a seminar dealing Klein instructions to lay the form 4.5 from the pre- with certain aspects of global change. We’ve all ceding as well as the succeeding stations underneath known for some time that the key to the interannual the one being worked on and use the observational and decadal changes in our weather is the oceans. points from those stations as a guide for interpolat- We’ve known that in an abstract sense, and more ing. This was an early form of what we now call recently we think we are beginning to understand objective analysis, but the computer does the work the relationships. Certainly the success that we are for us these days. I hope that somewhere on this now having in relating the so-called Southern Os- campus a few form 4.5’s still exist, along with the cillation to El Nifio events on the west coasts ofboth detailed Klein instructions, including, as I recall, a North and South America gives us hope that this shift in the hardness of the pencil depending on the complex interaction can be unraveled further. quantity being plotted. We also are beginning to understand the role of In addition to all that physical and chemical data the oceans in global warming, triggered by the in- carefully observed and recorded, there is the won- crease in such greenhouse gases as atmospheric car- derful biological collection of CalCOFI - the fish bon dioxide. One can perhaps best see the ocean role eggs and larvae, phytoplankton, zooplankton, and by examining two relatively recent models of how the fish, all carefully collected, carefully preserved, and how much the earth warms as the greenhouse and carefully archived. gases increase. One model, out of the United King- At a time of increasing concern about global dom, treats the ocean more or less as a boundary change, at a time when the prospects are increasing condition. It shows the temperature of the earth that we will soon be able to predict yearly and de- warming up less in. the tropics, but increasingly as cadal changes in our atmospheric and oceanic cli- one moves poleward. The heating is more or less mate, and at a time when we are looking for symmetrical about the equator. The second model biological and chemical trends over time, many are uses a three-dimensional ocean and is generated by scrambling to find long-term data sets upon which Suko Manabe and colleagues at NOAA’s Geophysi- to test ideas, to look for small signals hidden in a cal Fluid Dynamics Laboratory in Princeton. Like large noise level of what must still be treated as ran- the U.K. model, it indicates limited heating in the dom variability. The CalCOFI data set is almost tropics, and increased heating with latitude. But un- unique. Certainly there is little like it for the Pacific, like the U.K. model, it indicates that almost all the where our ideas about atmospheric ocean coupling heating is in the Northern Hemisphere; the Antarc- are more advanced. And there are few, if any, long tic Circumpolar Current absorbs the heat from the sets of biological samples where one can test trends Southern Hemisphere, with the result that there is in pesticides, for example-ideas not dreamed of almost no atmospheric heating at high latitudes in when CalCOFI began, using techniques not in- the south. vented at the time CalCOFI began. Like Fridtjof I’m not certain that anyone, including the model- Nansen’s International Hydrographic Bureau, ers, believes their predictions in any great detail, but which in retrospect was one of ICES’S greatest con- they do point to the importance of our understand- tributions to science during its first decade, so the ing the circulation of the ocean and its heat budget magnificent records of observations of the Califor- much better than we presently do. nia Current may be one of CalCOFI’s greatest con- We can certainly learn something about the grand tributions to future science during its first forty circulation patterns of the ocean from satellite obser- years. vations. From the pattern recognition that one can The care that Hans Klein and those who suc- observe by looking at the surface temperatures of ceeded him have given to the collection and analysis the ocean, and from satellite altimeters, one can of data ensures their quality, something that one can- measure the shape of the sea surface and in turn not always guarantee with some other long-term derive the equivalent of surface pressure maps of the data sets when attempting to use them for attacking ocean. But we cannot “sound” the ocean from sat- problems not originally contemplated in the exper- ellites as we can the atmosphere. We will need in situ imental design. CalCOFI is a wonderful program; it measurements of one kind or another if we are truly has a unique set of records, which can only become to understand the circulation and the heat budget. more valuable as they are extended in time. The data The CalCOFI data set provides us with forty years

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of monitoring one small but essential piece of the nate that those who began this program have con- ocean circulation. I expect that studying that data set tinued it. Having been party to a number ofjoint will play an important role as we design our total efforts, 1 can only assume that the cooperative ar- ocean monitoring system. Whether that system fol- rangements between this university, the state, and lows the CalCOFI form is not so important, but I the federal government have not always been easy. expect that the interannual and decadal variations That you have succeeded as you have is a tribute to one can find in the California Current as indicated the patience, the tolerance, and the statesmanship of by the CalCOFI data can be used as a guide. many. California, this nation, and the world are fortu- Congratulations, and happy birthday.

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PANEL DISCUSSION WHAT SOCIETY NEEDS FROM OCEAN SCIENTISTS IN PREPARING FOR GLOBAL CHANGE

Michael Mullin: We have arranged the afternoon some of President Bush’s key environmental advi- session as a discussion in which the panelists will sors. The scientists from various disciplines painted have an opportunity to make opening statements a very bleak picture of the effects of rapid urbaniza- from their individual perspectives about science and tion: deteriorating air quality, ever-increasing water policy and the issues that will be facing society in pollution, rapidly accelerating deforestation. global change. Among other depressing statistics, they under- The moderator of the discussion is Robert Sul- scored that society is destroying forest land at the nick, executive director of American Oceans Cam- alarming rate of one acre per second. The U. S. pro- paign. In addition to giving his own opinion, he will duction of synthetic organic chemicals has gone pick out points of agreement or contrast and guide from zero to over 225 billion pounds per year in the the discussion after each member has had a chance last 75 years. And of course the world’s fossil fuel to speak. We will, at that point, take written ques- use has soared. Chlorofluorocarbon emissions, tions from the floor. which were almost nonexistent before World War 11, are doubling every decade. Robert Sulnick: First let me thank Scripps for giv- The scientists agreed that we could attack some of ing me the honor of being the moderator for this these problems if we imposed a two-dollar addi- session. I am truly impressed by the quality of the tional tax on gasoline. That would reduce the use of panel and feel quite humbled by it. fossil fuel and thereby reduce these harmful emis- I want to briefly introduce this wonderful group. sions. But it’s interesting to note that not one of the On my immediate right is the Honorable Byron members of Congress who were present - and they Sher, assemblyman for the Palo Alto district in the included some of the best environmentalists in Con- state legislature. Next to Byron is the director of gress - stepped forward and volunteered to intro- Scripps, Professor Ed Frieman. Next to Ed is Bert duce that legislation. Politicians simply do not want Larkins, who is a fisheries biologist and now the to tell their constituents that solutions to serious en- executive director of the Alaska Factory Trawlers vironmental problems require increases in taxes or Association. On my immediate left is Professor dramatic changes in lifestyle. John McGowan, professor of oceanography at That is not to say, however, that the voters cannot Scripps. Next to John is Boyce Thorne Miller, a be aroused to approve dramatic programs to attack marine scientist from the Oceanic Society, recently environmental problems. As you will remember, in merged with Friends of the Earth. On my far left is 1986, sensing that the voters were deeply concerned Professor Harry Scheiber, UC Berkeley law profes- about toxic contamination of the drinking water, sor and historian. and believing that the state legislature and governor We will hear from Byron Sher first. would not adequately address the problem, several environmental groups qualified Proposition 65 for Byron Sher: Thank you very much. It’s a great the ballot. In the November election the voters, pleasure to be here. I appreciate the opportunity to while returning a very conservative, probusiness participate in this important panel. Our topic is, governor to office, also overwhelmingly approved what society needs from ocean scientists in prepar- Proposition 65, which, among other things, estab- ing for global change. My special perspective is, lished strict new prohibitions on toxic discharges what politicians need from science, although some into the state’s groundwater and surface waters. would argue that politicians should not be consid- Interestingly enough, for the first time ever, the ered a part of society, or at least civilized society. scientific community was drawn directly into that A few months ago the Smithsonian Institution election campaign. Experts in toxicology were used held a two-day conference in Washington on global both by supporters and opponents of the initiative environmental problems. Participants included to bolster their respective cases. One prominent UC some of the country’s most eminent scientists, sev- Berkeley professor, whom I shouldn’t name but eral prominent senators and congresspeople, and will - Dr. Bruce Ames - was pressed into service SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990

by the opponents of Proposition 65, which he stated warnings are predictive. Scientists are not abso- was environmental overkill. He appeared at public lutely certain that these changes will occur, and forums and urged voters to defeat the initiative. And many, or some, profess not to know whether the indeed his so-called peanut butter argument (eating proposed solutions will work. peanut butter poses a greater cancer risk than ingest- Secondly, the public does not yet perceive the ing drinking water containing minute levels of toxic physical effects of global warming, and conse- substances) became legendary during the campaign quently science has not convinced the public of the and was really the cornerstone of the business com- need for action. munity’s opposition. Third, many of the solutions offered by science Now that Proposition 65 has become law, scien- are perceived to be extreme or impractical. Reduc- tists continue to be involved in implementing it. ing greenhouse gas emissions seems to require enor- Under its provisions a scientific review panel has mous sacrifice on the part of the public. been set up and is required by law to determine And finally, other immediate concerns such as which chemicals should be listed as carcinogens or housing, AIDS, health care, and education are ab- teratogens. This panel has routinely been at the ten- sorbing all of the state’s limited resources. The pub- ter of heated controversies between elected officials lic is unwilling to divert funds from these important on both sides, and its determinations not to list cer- programs to attack an uncertain global warming. tain chemicals have even been challenged success- To return to the question before the panel: What fully in court. answers does society need from science in preparing My point is that science and scientists are being for global change? Well, here’s the challenge. Sci- drawn more and more into the political arena. ence needs to provide the public and elected officials Elected officials and society in general are demand- with precise, accurate information on the of ing hard-and-fast answers to complex problems. the problem, and a range of realistic solutions that But science often cannot provide such answers. And can be implemented within a reasonable time. State indeed, in those instances when science does pro- and federal governments will most likely not be pose a set of solutions, such as at the Smithsonian moved to action on global climate change in any conference, we are often told that those solutions are meaningful way until proposals of this nature are politically impractical. made. Global warming is one ofthe most vivid examples I believe there is little chance that significant of the dilemma created as science and political deci- amounts of California state revenues will be devoted sion making come together. The tremendous im- to broad-based programs to address the threat of pacts that scientists have predicted from global global climate change unless we get something climatic change demand responses. But so far Cali- that’s the political equivalent of the recent earth- fornia is doing little either to reduce its contribution quake. In fact, in this past session of the legislature, to global warming or to prepare for the effects that the governor vetoed a couple of modest bills that are predicted. would have inventoried greenhouse gas emissions in For example, the Department of General Ser- California and would have made a beginning on re- vices, the agency that oversees new construction of ducing those emissions. state buildings, is not considering global warming So science must become part of the education pro- effects when siting or designing new state facilities. cess. Unless the public is well informed and de- The Water Resources Control Board, which has ju- mands action from its politicians, there will be no risdiction over the groundwater and surface waters political will to address problems like global of the state, has told us that it does not intend to warming. address changes in runoff patterns resulting from In summary, there is a gap between the answers global warming in its current hearings to determine to problems offered by science and the solutions ac- the allocation of waters that flow into the Sacra- ceptable to the public and therefore to politicians. mento River Delta and San Francisco Bay. On the That gap needs to be narrowed. Conferences like basis of this experience, we can justifiably conclude this one and the media attention that is paid to it are that these agencies, and indeed state government as an important part of the process. a whole, are ignoring science. There are several reasons why scientific warnings Edward Frieman: I’m here under false pretenses; about the effects of global climate change have not you’ll have to pretend that I’m Bill Frazer, senior generated much response from political bodies, or a vice president of academic affairs, who was sup- demand for action by the public at large. First, these posed to be on this panel and is otherwise occupied

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with earthquake activities. I’ll base my remarks on A second initiative already under way is a joint the paper that he sent down. UC Davis-Lawrence Livermore-Lawrence Berke- He said, “It’s a great personal pleasure for me to ley Lab effort, funded partially by INCOR and par- extend congratulations on the fortieth anniversary tially by the Department of Energy, to conduct a of the CalCOFI program.” series of workshops on global greenhouse effect. He had been asked to provide an overview of the These workshops were designed to elicit the views university’s activities related to global change. The of international and national global warming ex- Global Change Advisory Group held its first meet- perts on problems of climate change. The first con- ing here at SI0just two days ago. The group con- ference took place in July, and was aimed at iden- sists of scientists from the campuses and the three tifying information needed to improve climate Department of Energy laboratories -Lawrence models and climate projections. The second confer- Berkeley, Livermore, and Los Alamos -that are run ence, held in September, focused on options for re- by the University of California. The group was con- ducing emissions of CO,. The third workshop, just vened to advise the office of the president on how to a few days ago, brought together a small group of build upon the already significant research being policy planners and researchers from fourteen Pa- conducted throughout the university on this very cific Rim countries to examine the causes of climate important topic. change and its implications. The group’s specific charge is to suggest means There is a third, very major, initiative under way, by which the university administration can most in which the university will participate. It’s called effectively support the design and functioning of a the National Institute on Global Environmental coordinated systemwide research effort on global Change and is just in its formative stages. This is a change, and to recommend elements that might multiuniversity effort involving the University of constitute such a research effort. California, Tulane, Indiana, and Harvard. It is writ- Five options were presented to the group for dis- ten into legislation introduced by Congressman cussion. These came out of a previous small rump Fazio and calls for $6 million in this first year and meeting we had over two months ago. The options $10 million thereafter. (This has caused some raised included establishing a research center focusing on eyebrows.) one or more of the nationally identified high-prior- Those are the three major activities under way. ity research fields; creating a research center without The charge to our group was clear. Two days ago walls to address one or more high-priority research we examined these various issues, and I guess it’s fields; expanding the INCOR effort to develop a fair to say that we are in the very formative stages. coupled ocean-atmosphere model, and you heard One of the first things we did was to try to get an something about that this morning from John inventory of what’s going on throughout all the lab- Knauss (I’ll come back to this in a little bit); creating oratories and campuses in the University of Califor- a global change minigrant program; and establish- nia. We tried to map that into the Committee of ing an international clearing house for information Earth Sciences’ famous document on U.S. global on research activities and data about global change. change research categories, which lists things in Presently there is a special project organized un- seven major chunks: climate and hydrological sys- der the UC Institutional Collaborative Research tems, biogeochemical dynamics, ecological sys- Program (INCOR), which was started in October tems and dynamics, system history, human inter- of 1988. We at Scripps, along with researchers at actions, solid earth processes, and solar influences. Lawrence Livermore and Los Alamos, won one The first result, which perhaps wasn’t surprising, competitive program with a project to understand was that there isn’t a one-to-one mapping; there is exactly how pollution and other factors are changing much more going on in the University of California the world’s climate. All three of us have supercom- that can be labeled under the rubric of global puter facilities and programs in climate research. change, perhaps with a small g and a small c. In The project focuses on developing a coupled ocean- particular, this White House document doesn’t atmosphere model for global change. The Scripps really address issues of mitigation. researchers bring their expertise on ocean modeling, In our deliberations we examined three possible Los Alamos its expertise in areas of atmospheric sci- ways to move ahead. One was to expand the collab- ence, and Lawrence Livermore the results of a dec- oration that now exists between Scripps, Livermore, ade of research on modeling the global climate. The and Los Alamos to other campuses. Two of them first report is due this December. The total cost of have major programs that would be a natural match: this four-year project will be $800,000. the atmospheric group at UCLA, and Sherry Row- SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990

land’s outstanding group at UC Irvine. Perhaps something about their reproduction. In fact we there will be others in the future. bought the Miller Freeman, brand new, the first U.S. The second point that was made was that it would factory trawler research vessel. I think that it was on be a good expenditure of a small amount of money its second cruise when we came down here to try to to have a number of us talking to each other. We find spawning hake. really don’t know from one campus to another We knew from your information that they what’s going on. The totality of the research going spawned somewhere around here, probably from on throughout the UC system, as addressed to this San Francisco to the southern end of Baja California particular problem, is just enormous. Some people in some years, sometimes out as far as anchovies made the statement-it’s hard to prove that it’s spawn- probably 150-200 miles. They were very true - that it’s perhaps the major powerhouse in the patchy some years. We had some new trawl gear, United States. and we wanted to get our hands on fecund hake. We The last thing we examined was the issue of arranged through Paul to have help from one or two whether it makes sense at this point to somehow of the ships that were out doing the CalCOFI sur- join forces across all the campuses for a new major vey. Overnight they sorted their plankton samples initiative. We’re still in the process of exploring this. from the previous day and looked for hake larvae or eggs. We started getting nightly reports that “Yes, Bert Larkins: Even though I spent the first twenty we’re getting more of them and they’re getting years ofmy career as a fishery biologist in the North- younger, and we’re heading west on such and such a west Fisheries Center of the National Marine Fish- trackline.” We’d run another forty miles west of eries Service in Seattle, I did get directly involved them and drag our net around and look at our echo in some of the information that CalCOFI was sounder. It took us about one week to find our first collecting. spawning concentration of hake. They were not Many will remember that about 1965 the Soviet where most people thought they would have been; fleet came sailing around the corner and parked right they were farther offshore and farther south that off the middle of Washington and Oregon and began year, as I recall. We would probably have spent our taking away 200,000 or 300,000 tons of hake a year. entire month looking for the first one if it hadn’t In its infinite wisdom our government thought that been for this very simple colleague-to-colleague we ought to know something about these critters type of arrangement. that the Soviets seemed to know so much about. So One thing I want to mention about what we’re we instituted a groundfish research program at the going to need from scientists in the future is to at Seattle lab, and I was its first director. least talk to each other and make sure that the infor- The Northwest rather prided itself in being way mation any of us has gets into the hands of our col- out in front of fishery research. It had the University leagues and of the public. of Washington and Oregon State University and the The organization I represent, as of two weeks University of British Columbia and two or three ago, is the Alaska Factory Trawlers Association. It others. But when we looked around for information is made up of about forty-five vessels, and has be- that might indicate something about what went on come unbelievably powerful in the last three or four off our coast, there was zip. years. There were three or four small factory trawl- So we got on a mailing list for CalCOFI infor- ers in 1980; now there are forty-five. Some of these mation. We did know enough about hake to at least are small, headed and gutted boats; they have a mod- surmise that they spawned down here somewhere. est need for high-value fish during the year. Other In those days the National Marine Fisheries Service boats represent $30 million and $40 million invest- was like a university; we had our little turf battles - ments; they have not just the capacity, but also the Northwest came down so far and Southwest went financial requirement for about 60,000 tons of pol- up so far, and never the twain would meet. I have lock a year. They can’t make their payments without kiddingly said that those were the days when Paul 60,000 tons; if they catch more than that they start Smith and I would talk to each other from phone making some money. Fifteen more of these big booths - we didn’t dare do it from the office. But ships, which make surimi and fillets, are on the ways nevertheless, we found out something about what around the world and will be in the fleet in another they knew about hake. year. So twenty years ago - halfway through your his- Right now we have a capacity - a necessity - for tory- we were able to learn something about these about a million tons of groundfish a year. Most of , something about their migration paths, this is pollock. Next year, or two years from now,

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we’ll probably need two million tons. The scientists mental to shellfish but was just wonderful for fin- tell us that pollock is probably the biggest single fish. Again, I don’t think anybody knows for sure. groundfish resource in the world now. It has an al- What’s happening now? I’m jumping around a lowable biological catch about 1.2 or 1.3 million bit, but I’m trying to relate some of the man-made metric tons in the Bering Sea and another 60,000 to problems that are going to occur in the fishing in- 150,000 metric tons in the Gulf of Alaska. This fleet dustry, particularly in the North Pacific. How they now has the capacity to take all of that. (There are get mixed up with natural events is anybody’s guess. no foreigners fishing this any more.) In another year It’s probably not going to be for the better when we we will have one and a half times the capacity for hit a natural downturn combined with all of the taking it. pressure that’s out there now. We had our sardine situation ten years ago in The fleet that I represent has now a billion-dollar Alaska. The big fishery at that time was king crab - asset value. As I mentioned, some of these boats are weight 10 pounds each. The ex-vessel price in $30 million investments, and there are others com- Alaska is now about $5 a pound. In 1980 the record ing on line that are 40, 45, and $50 million invest- landings that had been peaking every year before ments. Most of these boats have twenty-year mort- that were something like 80 million pounds from the gages on them; no longer is it a six- or seven-year Bering Sea. The next year the landings dropped in mortgage. half, the next in half, the next year in half. Now the As your experience down here would indicate, landings are maybe between 10 and 20 million I’m sure, fishermen are generally very good theoret- pounds. And this is after several years of some ical conservationists. They understand. And they rebuilding. mean it when they say, “I want my son to be able to Nobody quite knows why it happened. Was it a do this. I don’t want to overexploit. I want to make natural occurrence? Was it ? I think the a living ten years from now, and I want my son and best judgment is that it was neither; it was probably grandson to do it.” But then the bank says, “You both, very much like some people would guess owe me $lOO,OOO.” Well, suddenly they have to about your sardine situation. King crabs were in a catch as many fish as they can, very quickly, to make sort of natural decline; the fishery was very intense; this year’s payments. This is in spades now, in the and before people could get a handle on the decline, industry I represent. There are debt services that are that very powerful fishery drove it a little further unbelievable- $10,000 a day on some ofthese boats. down than nature might have taken it, and got it Their pro formas count on 270 days fishing. down there a little faster than nature might have. The allowable catch is rather conservative now, These are long-lived animals; they don’t mature un- because up to this point we had foreigners to boot til they are 6, 7, or 8 years old, when they enter the out if there was any concern. But now the foreigners commercial fishery. They live to be 15 or so. So they are all gone, and we’re competing among ourselves. are not like shrimp. Not only is there concern on the part of vessel own- At the same time, one species of Tanner crabs, ers that they need more fish to make their pay- which was not being terribly heavily exploited, also ments - which is the case, and it’s going to get even took a downturn. Most of the shrimp in both the worse as more of these boats come on line. What this Bering Sea and the Gulf of Alaska virtually dis- is going to do is put political pressure on manage- appeared. ment councils, on the government. John Knauss is Also about that time, and it took us two or three going to see a lot of this as the politics overflows years to figure this out, most of the finfish-like from the councils into Washington. pollock and cod - expanded to historic highs and are What we’re going to need is scientists who under- still there. These highs even exceed those of some of stand what’s going on in a multispecies, very com- the anecdotal reports from the turn of the century, plex, physical-chemical environment. They need to when the groundfish fishery was pursued by sailing understand the relationships of both man-made and schooners and dories. I’ve heard some people say naturally induced changes in relative abundance of that what happened in the Bering Sea, probably competitors, predators, etc. We don’t know very starting in the late 1970s, might have had some sort much about this now, certainly not up north, where of relationship, maybe an inverse one, to an earlier we are. Is it good to catch cod, because they proba- El Nifio down here that had some effect, not really bly eat king crab, even though you catch some king well understood, up in the Bering Sea. Others have crab while you’re trawling? said that for some reason a lens of cold water formed What we are going to particularly need from folks over the bottom of the Bering Sea and was detri- like you are scientists who are willing to get up, SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990

work in the fishbowl, and learn to speak lay talk so change will be, but almost everyone agrees that they councilmembers who have their hearts in the right will not be benign. place but don’t understand the jargon can under- We don’t know much about the rest of the global stand you. And then we need you to be willing to conditions or what we’ve done to them, but I sus- defend your conclusions. My experience has been pect that maybe other such large-scale changes have that out of 130 people at the Seattle laboratory, there gone on. But since we simply haven’t monitored were 6 of us who wanted to do this kind of thing; them the way we’ve monitored the CO, question, we wanted to be management biologists rather than we are unaware of them. We can be certain, how- research biologists. It’s been very rewarding for ever, that these, over time, will become more and some of us. We need a lot more people like that. more evident. So all of the marvelous progress we’ve made since John McGowan: Since I’ve had long experience World War I1 is now threatened in a new way. We with the CalCOFI program, I assume my role on must understand the magnitude of this threat. We this panel is to serve as some sort of a sea-truth absolutely must better understand what’s happening referee, or perhaps a dull scientific answer man. to us. While I’m willing, and may even be capable of serv- Because the oceans are so large and such a big part ing in that role, I have something to say as a private of the world, and because they serve as a sink for citizen. many pollutants, especially CO,, it’s crucial that we It seems to me that there have been some very understand how they function and how their biota important changes taking place in the world, espe- responds to changes. We must understand how to cially lately, and there is really much cause for opti- measure change and its direction and magnitude. We mism. The threat of nuclear war or even large-scale must understand its consequences. conventional war is greatly diminished; human The CalCOFI program - with all ofits faults, and rights and even human welfare are now being seri- I understand many of them - was designed, in the ously considered in many places; there have been first place, for the purpose of studying the magni- relatively peaceful, popular, revolutions, major tude and scale of environmental change. Although ones, as in Poland, Hungary, and other places. De- we’ve made a lot of mistakes, many of which only mocracy has become a fashionable word once again. those of us who work in the program fully under- We have now institutionalized the prevention of the stand, we’ve made a lot of correct decisions as well, great killer diseases such as smallpox and cholera. and have learned much. We understand now how to We can grow enough food to feed the world, al- go about studying the large-scale problem of though we don’t distribute it very well as yet, and change, and it is the CalCOFI data that will help us even the fine arts are flourishing. It’s beginning to do so. We’re very fortunate to have this marvelous look as though humankind has learned how to deal data set, for it can serve as a basis from which to with some of the great traditional adversities and proceed and as a template for further study. To quote afflictions that have been with us for so long. someone from outside the system: “That data set, But while our leaders and policymakers and poli- right now, is a national treasure.” There is, in my ticians have been making this remarkable progress, view of things, room for optimism; we in fact an entirely new and unprecedented challenge has started to study the problem of change forty years arisen, one with which we have not had any substan- ago, and we now know how to go about it. This is tial, collective experience. I’m talking about our re- an enormous advantage. lationship with nature. This has become a critical issue, and it may become acute. The problem is so Boyce Thorne Miller: Global environmental large and so complex that it can hardly be stated change, as we are discussing it today, begins with coherently. In the next twenty years or so we’ll hear humans and ends with humans. In between that be- much about it. ginning and end there will continue to be significant It differs from previous problems due to our dis- impact on the land, the oceans, the atmosphere, and ruption of the natural order of things in that its scale all the life that knows this planet as home. Bill is much, much larger, even global, and it is, there- McKibben, in his powerful new book, has noted fore, much more complicated. Although we are that this era of human-induced global change marks short of much factual data, we know for sure that we the “end of nature” - the title of his book. have managed to change the atmosphere itself. What No longer are humans responding to the forces of is not so clear is what the consequences of that nature; now nature is responding to the forces of

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human culture. We are dealing with a nature molded ment agencies to allocate more funds for such work, by human beings. Because of this, science is also possibly even from that seemingly untouchable changing. Scientists studying natural phenomena ocean of funds now set aside for defense. The envi- are now more often measuring and predicting na- ronment, after all, is an issue of global defense. ture’s response to human manipulation of the envi- I may need to work a little harder to convince ronment. And they are called upon to distinguish some of you scientists to participate in environmen- natural variations from human-caused variations, tal and management decisions at local, state, federal, though the two are often inextricably meshed. and international levels. Ocean scientists are partic- There are two human factors driving this over- ularly important because the coastal zone is quickly whelming influence we have on the natural world: becoming the front line - the place where the envi- explosive population growth and explosive eco- ronmental battles will be first and biggest, because nomic development. Unless these are curtailed, we most of the pressures from population and economic cannot hope to slow the rate of global change. And development are focused on the coastal zone. Ever- it is that unprecedented rate that has led many sci- increasing stresses are placed on the coastal environ- entists to worry about the ability of species and eco- ment as a result of industrialization, urbanization, systems to adapt. There are many ways our indi- residential and tourism development, waste dis- vidual lives and the structure of our societies will charge, dredging, poison runoff, and overfishing. have to change if we are to succeed in slowing global Also, agriculture, forestry, and mining in interior as change. But that is a discussion for another forum. well as coastal regions create runoff that eventually Today we have been asked to speak about ocean enters coastal waters. scientists and what they can do to help prepare for The threats to the coastal zone from all this human global change - and, I would like to add to that, to activity include pollution from overenrichment, help moderate global change. I would suggest to toxics, and debris; habitat destruction, particularly scientists three contributions that you can make. on the coastline; and overfishing. Many coastal eco- You can do basic and applied research relevant to systems are already impoverished. And even deep global change; you can aggressively involve your- ocean waters are not immune from the impact. Our selves in environmental policy and management de- living marine resources are being jeopardized on a cisions; and you can take a strong public stand on global scale. environmental issues. Now, how does this apply Problems created by multiple usage of a fluid en- specifically to ocean scientists? vironment that cannot be compartmentalized and I have no doubt that research is the easiest for me does not honor political boundaries are complex. to convince you to do. We need more information The solutions are also complex and must be innova- about the ocean’s role in climate change, in the ox- tive. Sound decisions about coastal policy and man- ygen and carbon cycles, and in other geochemical agement and how best to regulate human activities cycles. We need to learn more about how the ocean in the coastal zone require scientific and technical may influence and respond to global warming. We expertise. Decision makers often don’t have this ex- need to know more about the diversity of life forms pertise. So it is imperative that marine scientists get and biological processes in ocean ecosystems -how deeply involved in the process. they compare to terrestrial ecosystems, how they I am asking you all to participate - locally, nation- will respond to global environmental change, and ally, and internationally. One word of caution, how- on what time and space scales these responses will ever: don’t expect this involvement to be a particu- occur. Fisheries biologists need to learn more about larly rewarding experience. Don’t expect to come the effects of pollution and harvesting on the health away each time feeling that politicians and environ- and abundance of fish and shellfish populations. We mental managers will act on what you said, even if also see a need for more well-designed, long-term they listened. It is not an ego trip, and it requires environmental monitoring programs in coastal and perseverance. Keep going back with your message. ocean ecosystems worldwide, similar to the Cal- And you will have to learn to simplify your mes- COFI program. But ocean scientists are eager to do sages as much as possible. Those making the deci- this work. I don’t have to convince you to do basic sions often find it difficult to convert very complex and applied research and monitoring. scientific information into policy or regulation or Better that I sit here as a representative of the management practice. So it is up to you to bridge environmental community and tell you that we are that gap. And we of the environmental community working hard to try to get Congress and govern- can help also.

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Let me give you some examples of what you can But for environmental decision making, it is impor- do. A very simple thing you can do if you are at tant to take strong action on strong scientific likeli- Scripps is to march down to your aquarium and tell hoods. We can’t wait for absolute proof before we them to stop selling shells plundered from Philip- act, because the proof comes after the damage is pine reefs. Another thing you can do, on a larger done. scale, is to involve yourself in the EPA decision about sites for dredge disposal off the coast of Cali- Harry Scheiber: As a person trained in economic fornia. The EPA seems to feel that there are no data. history and legal history, doing a considerable Knock on their door, or make a telephone call and amount of teaching on law and technology and on tell them about the CalCOFI program. the law of the sea, I found this a daunting assign- The secondary treatment issue is a big one, a con- ment. I think that in some ways we who are trying troversial one. Many scientists have said that off the to see the proper linkages between matters of policy coast of California you don’t have to have secondary and law must be more responsive to environmental treatment, or at least that’s not the best use of funds. needs and crises. Let’s look at it in a slightly more innovative way. It We’re in much the same state as the scientists of may be that pretreatment will have more of an effect CalCOFI were forty years ago - a state of great per- than secondary treatment. However, both of these plexity. If you were to draw a historical time line, as together would be even more useful. We cannot af- I’ve been seriously playing with in the last few years, ford to waste our wastes by throwing them into the of the relationship of changes in what we today call ocean. They should be put back on land and used. fisheries oceanography and international policy and Get the toxics out, yes, and then use the organic law, I think there would be, from World War I1 to matter to fertilize our fields. the present, working backwards, the following mo- Finally you can work on an international level. ments of really fundamental change and innova- International cooperation of scientists, as you have tion - turning points. here in the CalCOFI program, is admirable and One, I think, will turn out to be the current dis- should be encouraged. The regional sea programs cussion of global change, which really started in a offer another opportunity for that. International serious way two years ago. lending agencies need some advice. They believe Going back a considerable period, we’d go to the that economic development is what all developing 1972 Stockholm Conference, which brought to- countries should be aiming toward. So the scientists gether worldwide United Nations representatives to in those countries -in Mexico and in other devel- discuss the problem of environmental crisis. This oping countries -need to get the message across to was certainly another of those watershed moments their governments and to the lending agencies that when our thinking was fundamentally changed there is now a new goal-sustainable use of re- around a really significant problem. sources. Going back again - a leap of fourteen years - we The last suggestion I have is that you scientists come to 1958, when there were conferences on the take a strong public stand on environmental issues. law of the sea in Geneva and then Rome. They were This is not for everyone. You have to tread a fine line the beginning of the law of the sea movement, between credibility and effectiveness. So I ask those which has culminated thirty years later. At those of you who don’t choose to go public to be tolerant meetings we began to discuss the possibility of some of those who do. Don’t criticize them for being kind of convention that would bring the nations of what may seem to you too simplistic. Remember the world to agreement on the regulation and con- that the public and policymakers do not quite know servation of the sea’s living resources. There was a how to deal with all the words of caution and con- fundamental transforming effect, not only on those ditions that you put into your scientific conclusions who tried to create a law of the sea - a long, diffi- when presenting them to your peers. And they often cult, somewhat frustrating and not altogether happy use the uncertainty to further personal, institu- process, which has culminated in the current law of tional, and political goals. the sea convention. At the time, the introduction of For example, the World Bank recently declared the sustained yield concept, of the idea that the goal global warming a lionissue in its lending policies should be maximum sustained yield of ocean re- because of the scientific arguments over the magni- sources over time, crystallized thinking in a way that tude of what the impact will be. For the purposes of had not been done before. solid scientific research, it is important to be aware In 1952, six years earlier, came the first of the of the weak links and the possible sources of error. postwar treaties of the Pacific, which Japan signed

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after its sovereignty was restored. This treaty in- its parts if they could be brought together. Some- cluded for the first time in the postwar era the con- thing that hasn’t been mentioned, but that I think is cept of maximum sustained yield. Japan, the United worth underlining, is that industry supported it States, and Canada agreed with respect to salmon, from the start. The industry had a specific problem halibut, and herring that they would maintain the and went to the scientists for the solution. goal of maximum sustained yield. From that came This brings me to the second point, which is that the very controversial abstention concept. the scientists’ response was extremely creative and, These, I think, are the major turning points - the in a fundamental way, “subversive” from the begin- watershed events when our thinking really crystal- ning of CalCOFI - California Cooperative Oceanic lized in new ways. Fisheries Investigations. When this group first But first, on this time line, really has to be Cal- formed, the Marine Research Committee, which COFI. You may be startled to think that your being granted money for collaborative research, said, “We out on the edge of the continent worrying about the have to have a name for it.” So John Marr of the sardine and the California Current is such an event, federal service immediately provided a name that but I don’t think we exaggerate. In this venture- would fit nicely into the federal hierarchy and table now forty years old, an extraordinary length of time of organization - the California Sardine Investiga- for such a venture to have survived and be still so tion. Frances Clark of the state agency responded vital - we have some lessons to be learned. Not just and said, “Let’s call it the Marine Resources Investi- about why things developed as they did later, be- gations. ” Roger Revelle came up with the California cause there are lines of continuity here, but also Cooperative Fisheries Investigations as a compro- about how things can be done in the future. I’m not mise. But of course it wasn’t a compromise at all. going to be able to talk about all of these; I just want From the outset it announced that CalCOFI wasn’t to suggest that this is another in the series of impor- just about sardines - it was also about fisheries in tant events in which thinking was crystallized on general. new lines. From the very beginning there was an ecological Some of the consequences were not at all unantic- vision as well. We think of it as something that ipated. There are two interesting elements that I’d evolved and was produced over time, and we asso- like to single out. One is the collaborative element ciate it with the late sixties. But in fact, in the very from the very beginning. The other is something first documents that were circulated - from that was discussed as a longer-term consequence and Scripps, from Frances Clark, and from Oscar Sette outgrowth of the beginnings, and that is ecological of the federal agency - the idea was inherent that vision - the ecosystemic approach to fishery prob- (with the new equipment and the new ships that lems. Roger Revelle had produced as a gift from the navy, The Marine Research Committee was approved to President Sproul’s shock, and with the new fund- by the legislature and funded from the beginning by ing) for the first time they could get answers to ques- a rather sizable tax on sardine landings. CalCOFI tions that had been plaguing them for twenty years, was a response to the sardine crisis, at the beginning. and which they alveady understood were the important There is a little two-year prehistory here, which questions. So in many ways the scientists’ vision hasn’t been mentioned. It was the formation of the prevailed here. Marine Research Committee, authorized by the leg- From the very beginning this vision was set forth; islature of California in the winter of 1947. The the scientists had an agenda right from the outset. committee was financed by an industry tax on the The lesson was not lost on the industry, which was landing of sardines and was really the progenitor of a little dismayed, but which continued to support CalCOFI. It was the response to a crisis. the investigations. Over time, people in the industry From the outset of the project there was a collab- and in the outside world supported the research, in orative intent, with the argument made that it considerable measure because of the efforts of a sci- would bring together for the first time the resources entist who was also a great organizer - Wilbert M. of the Scripps Institution of Oceanography and the Chapman, a very important force in the beginning, University of California generally; a federal first working out of the California Academy, then agency - the Fish and Wildlife Service; and the Cal- in the State Department, and later in the industry. ifornia Department of Fish and Game, in which This project was watched closely from the begin- Frances Clark had done many years of very impor- ning by ocean scientists in all nations. As John tant sardine research. From the beginning there was Knauss said this morning, it could be that its role as the vision that the sum would be much greater than a prototype was understood from the beginning.

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To wind up where I began, one of the really inter- explanation has to do with all of the non-point esting things about the documentation of the meet- source pollution that flows in from all the rivers that ings of the scientists who set up the protocol for the enter into the bay. 1952 Japanese-American-Canadian convention and I have been up on Puget Sound, where I’ve seen also many of the scientific papers that were pre- liver cancers in sole. And I’ve heard the explanation sented at the Geneva UN meeting, was that they that they come from the toxins that the industrial talked about the research done in the California Cur- complex puts into Puget Sound. rent as exemplary of what had to be done in order to And I’ve been in Boston Harbor and Deer Island, make a concept like maximum sustained yield con- where I’ve seen raw sewage in an overwhelming sistent with the best science. So the influence has display - emptied daily into Boston Harbor. That’s been very great over the years. a very serious insult to the integrity of that eco- The lesson, if you want to draw a lesson from it system. with regard to the role of science, is that a project I’ve been around the country and have heard hor- well founded organizationally and well conceived ror stories time and time again. In addition, I’m scientifically is going to have an integrity of its own given intellectual input. By next year 75 percent of and a vision of its own. And room has to be left for us will live within 50 miles of the coast, and we are that vision to be nurtured. And that has been done toxic-dependent as a society. We humans, gathering remarkably well indeed in the forty-year record of on the coast, are dependent upon toxics that inev- CalCOFI. itably, as a waste stream‘, make their way into that coastal zone, which includes estuaries, bays, and Robert Sulnick: Let me begin by saying that it’s coastal wetlands, and then kills the vitality of that obvious to me, an environmental lawyer for over coastal zone. twenty years, that global change is inevitable and I am trained to be an advocate, and I now instinc- that we will all have to be involved in it. If we are tively take all of that information and begin to cam- not involved the change could be so disastrous that paign with it. I do this without even thinking about the human species will no longer thrive or possibly it, as do all my colleagues. The campaign is aimed even survive. at the the public at large, the general population, to So my approach to this discussion is going to be seek a critical mass, and then at the decision makers somewhat practical. My assumption is that I’m ad- in Congress. Because we want that critical mass to dressing an audience made up mostly of scientists. I be translated into public policy. myself am an environmentalist and an activist. I Because I’ve been a lawyer for twenty-five years, would like to give you an insight into how the peo- I have learned that decisions will be made irrespec- ple that I work with - my side of the fence - think tive of who has the input, when a problem becomes about these problems. Because my premise is that large enough to be perceived by the political body. unless we come together, unite, and go forward, the It’s a reactive body. So we in the environmental planet is in very serious trouble. community are going about our business of trying When I think about the oceans, and when I talk to to create public opinion - critical mass (i.e., pres- people about the oceans, as I do all over the country, sure) - and then we are trying to translate that into I say this: “The oceans are the lungs of this planet. public policy. We will do that irrespective of the They provide us with 70 percent of our oxygen. ” I scientific input. don’t know that that’s absolutely true, but I’ve done The danger is that much of what we do, by defi- enough reading to know that I’m in the neighbor- nition and not by any conscious intent, is polemic. hood. And I also know that it grabs everybody’s It’s not meant to be polemic, but we work in such a attention, from people in the White House to people rampant atmosphere ofno resources, no money, and on Smith Island in the middle of Chesapeake Bay. no support system that we can only do the best we So I will continue to say it, unless people like you can. I don’t have five days to research anything, tell me, “YOUreally can’t say that, because it’s abso- normally. I’m either on the phone with the press or lutely not true. ” I’m in the field or I’m at a hearing. My schedule is I have been to Smith Island in Chesapeake Bay jammed. And again, I’m not talking about me as an and spent days with the crab fishery there. They individual but me the species. So we work with have shown me that from their point of view that what we have. fishery is disappearing. And when you look at the The danger is this: we move very quickly. And habitat through their eyes, it is disappearing. Their we will win our political battles because the public

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now wants us to. But in winning the battle, if the says that humankind has come to dominate the solution is ineffective, we may lose the ultimate planet, and we’ve carved our initials so deeply into struggle. the biosphere that we can no longer consider nature Your danger, it seems to me, since I have a back- to be separate and pristine. ground in social science and methodology, is that So I see some sort of a mini intellectual trend. I’m you can be restricted by two things: your methods, no historian, but my academic colleagues tell me which obviously you need, but which can restrict that this is a phenomenon that has been seen before you because of the time involved; and the money to as the end of a millenium approaches. finance your methods, which is probably the root of But rather than accept these pronouncements by the issue if we’re honest about it. Because money is these intellectuals and philosophers, I prefer my not being given to science, much less ocean science. own favorite philosopher - Yogi Berra, who said 95 And the next generation of ocean scientists is not percent of the experts in a certain field agree that getting in line, which is quite alarming. The money such and such is the case, and the other half believe is obviously going to military spending and has the opposite. It seems to me that is, in fact, the been for quite some time. situation we are in with global warming. However, we need to slow down, and you need to We do not, I’m afraid, have the unique footprint, speed up. Somewhere in the middle there’s a balance the scientific evidence to move ahead. You heard to be made. A dialogue must be created on how best from John Knauss this morning about major global to go forward. It’s no longer a question of should we models, one ofwhich says the Southern Hemisphere go forward together. The question is, how do we do does not change, another of which says it does it? Because we communicate in much different change. You will hear, if you choose to talk to Tim terms. We’re out there in the spotlight all the time Barnett, who has analyzed the data of Jim Hansen articulating, and you generally are not. I would not (who testified in Congress last year that the global expect that to be natural for you. You’re very cau- warming signal is updn us) and has printouts from tious and methodical and precise in what you say. Jim Hansen’s monstrous computer code which in- We do not have the time or, in our heads, the luxury dicate that even if you add no CO, the temperature to do that. So the question is, how do we blend those goes up for fifty years. So it seems to me we are, two things? unfortunately, left in the position ofhaving both feet It seems to me that’s what this panel has been firmly planted in midair. talking about and what we need to continue the dia- I guess I feel, as the director of a scientific institu- logue about. tion, that we as an institution are responding to the University of California’s motto, which is “Re- Frieman: A number of issues have been raised that search and Teaching and Public Service,” and we can are of vital interest to the scientific community, and serve the public by somehow trying to get at this I’d like to try to address a few of them wearing my vague notion of objective truth. director’s hat. I agree with my colleagues here. We have a very Let me start with the philosophical point of view. serious societal issue on our hands. And I have ab- I see a mini intellectual movement going on. I’ve solutely no problem at all with Scripps scientists seen the essay by Francis Fukuyama earlier this sum- speaking out on these issues as members of the pub- mer from the State Department policy planning staff lic. But I do think we have a responsibility not to do concerning the end of history. Fukuyama tells us it as an institution. We have an issue of credibility on that there is a triumph of capitalism and western our hands, and we can debate that for a long time. liberal democratic thought, and we’re condemned Let me turn to the second issue that has been to a boring and static future. raised. I look at the 1990 research plan for the United In the last week or two we’ve seen reports of con- States to tackle the fundamental issues we’re talking ferences on the end of science- that somehow we about. I look at the total expenditures for fiscal 1990: have lost our methods of dealing with objective real- $190 million. This is across all the agencies of the ity, followed by a report (in probably the same news- U.S. government. And I look at the fact that our paper) on exciting new discoveries from SLAC at total GNP across the world is $14 trillion, and I try Stanford and CERN in Europe on new particles that to argue, well, suppose the U.S. contribution of are the foundation of our universe. $200 million is just one-fifth of the total. I have no And I read Bill McKibben’s The End dNuture idea whether that’s true or not; I suspect that maybe when it was first published in The New Yorkev. He it’s wrong. But maybe the total amount of research

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we are expending throughout the world, trying to Professor Scheiber talked about watershed events cope with this problem is $1 billion out of a $14 in history, which, in my mind, is like reaching a trillion GNl? It is just ridiculous to assume that critical mass toward moving consciousness forward somehow we now have the science base to make in a given area. industrial decisions that will make a major change All of these points raise initial questions that I in a $14 trillion GNP would like to direct to each of you. I don’t know how to cope with this. It’s clearly a To you first, Assemblyman Sher: What if, in fact, major issue for all of us to somehow get the major science cannot make its interpretations and its find- research engine moving. We must do that, and we ings more accessible to the political process? How must convince our elected representatives to do it. do you in the political arena then reach out to the At the moment it seems to me that we are in a very scientists so that we can still have the input? serious situation in which we are trying to make vast Sher: First of all, let me say that as politicians we global decisions on the basis of an extremely poor don’t need scientists who are captured by people data base. who resist making changes, who are impacted by some of the regulations that we establish in reaction to problems. Unfortunately there are a lot of scien- Sulnick: I’d like to establish some ground rules for tists, as there are a lot of medical people and a lot of what I hope will be a dialogue among all of us. I’ll lawyers and other specialists, who are out there and synthesize what I heard everyone say. Then I would available. I remember a debate we had about so- like to direct some individual questions to the panel- called noncriteria air pollutants - not the things that ists, questions that I have been writing down as they cause smog but some other bad stuff. And we were were speaking. Then I would like you, the audience, told that our solution to a problem that we knew to involve yourselves in their responses. And then existed was based on bad science. There were expert we’ll move on to questions from the audience. witnesses for the industries that would have been As I listened to everybody talk, this is what I impacted by the regulation who told us that. The heard. (If I misquote anybody, by all means let that answer was, don’t do anything. be part of our dialogue when I’m finished.) First, I I think we’re going to see the same thing in the heard Assemblyman Sher say that it is important for global warming area. Only this week we saw Pres- scientists to make themselves more accessible to the ident Bush’s proposal for a new approach to pesti- political process, so that the political process can cides, and it contains some good recommendations make use of the science. to help the EPA respond more quickly. But at the Professor Frieman, when you gave your initial same time there is a big debate about whether the remarks, I understood that there is a large need now federal government should preempt efforts by the to study things on a global, systemic basis, maybe states to impose more stringent standards. The EPA even instead of on an individual scientific basis. representative I heard debate this said, “We set our From your last set of remarks I understood that this standards based on good science, and we don’t want institution as an institution ought not involve itself the states to [complicate the matter].” in the political debate, but that its individuals are I don’t think I’m answering your question, but I free to do so. wanted to get all these things off my chest anyway. What I heard Bert say was that it’s crucial to prac- Sulnick: Let me ask you this question: Let’s assume titioners who make their livings from the ocean re- that science could not give a precise, accurate solu- sources to have correct data so that their institutional tion to a political problem but would give a solution approach to making a living from the ocean will be that is imprecise from a political point of view and effective. not readily accessible to a political body. The way I Professor McGowan, when I listened to your el- view it, there will be one side in favor of social oquent remarks, I heard you say that the things we change and one side opposed to social change, at its have achieved since World War I1 are now being most simplistic level, in a legislative debate - those threatened and that we must approach the global who are going to push toxics reform and those who problem or lose all of the benefits that we’ve gained are going to favor the status quo. What I’m hearing in the last fifty to sixty years. you say is that whenever you have scientific input I heard Boyce say that it’s important for scientists that is not in favor of pushing toxics reform, the to get involved in the solutions to environmental opponents seize on it and say, “Well, see, there’s problems. really no reason to reform, and our industry

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shouldn’t have to spend millions of dollars on an concerned about preserving what we’re trying to imprecise solution. ” protect while the science is going on. I think the state Is there any way that you see, from your years of has an obligation to use tax money in significant experience in the legislature, to still involve science amounts to promote the scientific study of these in that debate and not give up the fight? Because problems. But what do we do in the meantime? what I’m hearing you say is that when science comes We’re told that tremendous impacts of global in and says, “Look, we really don’t know what the warming are coming. I would say we ought to at effects of global warming are going to be,” then least make a start on trying to respond. What are the everybody says, “Well, why should we spend bil- agencies that have jurisdiction over different areas, lions of dollars on global warming?” Even though like the Department of Water Resources, doing to intuitively people believe and know that there is pian for the potential impact of global warming? going to be a horrible effect of global warming. Whatever we do, we should do it in the context of How do you deal with that? existing programs, so that we’re promoting the un- Sher: I don’t think there is a way to do it. When derlying policies of those programs. For example, someone has convinced a member of the legislature there’s a good reason to cut down on carbon dioxide or even the governor that we have a serious problem for reasons other than global warming. So if we can that we ought to be addressing, and that issue comes bring the global change considerations in to help us up and we start to debate it . . . if there is this differ- do what we are already trying to do for other rea- ence of opinion (and I hear some scientists tell us we sons, we may have more success. have a new ice age coming), people who would be Sulnick: Would anybody on the panel like to corn- adversely affected by the proposed regulations will ment on this issue of science in the political process? seize on that difference of opinion as a reason to take From the moderator’s point of view, it’s a big issue, no action at all. because those of us who get involved in the political And what frequently happens is that the process process do so as advocates. We rarely get involved is long and complicated, with a scientific advisory objectively. And we’re looking for people to support panel. And nothing can be implemented until the our positions. That’s the contest. And of course proposal works its way through the deliberations of that’s not a scientist’s approach. And yet scientists the scientists. Meanwhile the problem is overtaking have an enormous impact on that process. us. Let me give an example of one that I worked on Larkins: I started to touch on this in my presenta- this last year. tion. A year or two ago at the University of Wash- As an environmental activist in the state legisla- ington I participated in a symposium that had to do ture, I’ve been trying to do something to protect with science and fisheries. I made a lot of my old what’s left ofthe ancient forests in California, partic- colleagues a little miffed - and one or two of them ularly the redwood forests, where there used to be 2 very pleased (the one or two who happened to share million acres and now we’re down to 100,000 acres. the same view). My training was in fisheries biol- (And almost 20,000 are owned by one company ogy. I worked my first ten years as a researcher. I that’s cutting them down in order to pay the debt on became very interested, on my own, in the applica- the junk bonds that were issued when they were tion of science to fishery management. This was taken over by a conglomerate.) What the industry back in the days before the National Marine Fisher- has proposed, and what has now passed, is a three- ies Service had any management authority. We were or four-year study of the ancient forest to determine a research institution. whether there really are ancient-forest-dependent We saw the Magnuson Fishery Conservation and species. But in the meantime I tried to get some Management Act coming along in the seventies; it constraints put on cutting down the forest while looked as though NOAA, which was fairly new at we’re studying whether we need it to preserve the that time, was going to have some management re- spotted owl and other ancient-forest-dependent crit- sponsibilities. There were a very small number of us ters. But we were unable to do that. The industry who just jumped at this opportunity. I guess we felt supported the study, which they will use to say, “We a little frustration. We had been doing science; we don’t need to do anything now because we’re mak- had some conclusions; and there was nobody to give ing the study. ” them to, nobody who was going to do anything We can’t have a in-house panel of scientists that with them. No one had the wherewithal to do it. we depend on who will inevitably direct us to the So there were a few of us who found ourselves in right thing to do. But I do think we have to be a different frame of mind. And first of all, as we’ve

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heard from many of the speakers today, it was ob- strongly presented science that might be available, vious that we had to learn to speak human, rather they usually make the very worst choice. than science. I don’t know how many of you are What all of this boils down to is part of what upset familiar with the council system that the Magnuson some of my colleagues a little: I see a need, way Act set up, but it’s the fishery management councils down in the science education programs, to start that advise the secretary of commerce, through Un- having people who are going to be in science under- der Secretary Knauss, about fishery regulation. stand that there may be two ways for their careers to They’ve become a little bit more than advisory. In progress. At least make them aware of these two effect, what they say goes, unless they’ve done avenues that they might follow as they get into their something blatantly illegal, and then the secretary careers. One is as a researcher; that’s finding knowl- of commerce may override them. edge for the sake of knowledge. The other is apply- The folks who are appointed by the political pro- ing knowledge. Somewhere, probably at the cess to these fishery management councils quite graduate level, certainly when one gets into a gov- often come from the industry. This is unique. A lot ernmental agency, there ought to be two distinct of them have vested interests in the fishing industry career tracks. Those who have the talent and the that they have been charged to give advice on man- wherewithal should be encouraged to get out, live aging. They’re not scientists, and they say, “Well, if in the fishbowl, translate science into lay talk, and the scientists can’t give us 100 percent assurance, get the word across to the policymakers. don’t shut us down.” McGowan: This issue of advocacy on the part of People in my association (I’ll probably get fired if scientists is a serious one. I’ve thought about it a lot there are any of them here) do that. Part of my job, because it has come up many times before. I’m very as I see it, is to advise them when to back off, so that leery of having scientists advise policymakers and they have something to be operating on five or ten politicians on solutions to some of our environmen- years from now. tal problems -not on the nature of the problems, In any case, this is a long way of saying that I see but on solutions. Because scientists’, or anyone’s, the federal service - the National Marine Fisheries advice is always value laden somehow. There’s a cer- Service - and the state agencies as being applied sci- tain amount of entrepreneurship involved, and self- ence agencies. This doesn’t mean they can’t do some promotion. Scientists, for better or for worse, often very fundamental research; I think they have to. But have a rather narrow view of the world. They come on the other hand, I think their job is to get their from a part of society that is remote from real world conclusions out to the public where they can be problems and concerns. I don’t think that their no- used. And by God, go out and do it as advocates to tions about solutions to environmental problems or make the policymakers understand what the scien- other serious problems are any better than anyone tific ground truths are, to the extent they can be else’s. In many cases they’re worse because of the articulated. narrow value judgments they put on things. A lot of us in fishery biology like to tell ourselves I think the judgments about solutions should that we’re worse off than most other scientists be- come from society, from people who have the infor- cause it’s such an inexact science; it’s almost an art. mation. It’s our job to present them with the infor- You put a little net down in the middle of an ocean mation. And the decision then is made at some other and get only a very small sample. I’m not sure we’re level. unique in that. But what I’ve tried to tell some of the Sulnick: Given that the decision will inevitably be young fellows that were working for me when I was based on value judgments, which are often subjec- still a fed is: You guys have to have the courage of tive, why should a scientist not offer his or hers? your convictions; you’re the experts on the biology McGowan: Because they represent a very small of these animals. The Magnuson Act requires use of segment of society, and a rather privileged segment the “best available scientific information” and that is of society at that. Their educational background is what you must provide. No “insufficient data” cop- rather specialized, and they often lack even a rudi- outs, but the best you can do with the data at hand mentary idea of human conditions. I’m not at all and your best scientific judgment. I’m not asking certain that they’ve got the best interests of society you to be biostitutes. But put the outside parameters in general at heart. (laughter) around it; tell them what your advice is as an expert. Scheiber: Just a footnote to that. At each of those Because if you don’t they are going to decide the watershed moments in the history of fisheries ocean- “science” by themselves. And when policymakers, ography over the last fifty years, the scientists did even those who mean well, don’t get the kind of take a very strong position on what needed to be

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done. For example, in the case of CalCOFI, there way we live, considering how many of us there are, really was an agenda. A few scientists had done exacerbates the problem. some brilliant work at the University of Washington Obviously the ten million peasants have to be fac- under William E Thompson, here at Scripps Insti- tored into ‘the solution. But you can still create a tution under Harald Sverdrup, and at other places solution that preserves the rain forest, that promotes including the California Department of Fish and the health of the planet that we’re all dependent Game under Frances Clark and Richard Croker. upon. And you don’t pit them against one another. Those people knew what had to be done, and they That’s a huge mistake, because no government can said what had to be done. turn its back on its people and still be in power. So The same was true of maximum sustained yield the people are part of the solution and need to be at the United Nations. Richard Van Cleve gave a factored in, and need to be part of the dialogue. very influential presentation that was based heavily Frieman: I have spent far too many hours and years on the work that had been done by CalCOFI scien- advising the government on one issue after another tists, who said, “This is what we need to do.” So to feel very sanguine about the prospect. We saw there have been moments when scientists have spo- what happened to the President’s Science Advisory ken out and have spoken out constructively. Committee when they recommended against devel- Listening to Dr. Frieman, I think there’s a differ- oping the supersonic transport because of environ- ence between those cases and this one, because there mental concerns. PSAC was then abolished, and it is -as he said so eloquently - a lack of an agreed- didn’t exist through the rest ofthe Ford Administra- upon agenda today on this larger question of global tion or the Carter Administration; then something warming. was put back together under the Reagan Admin- McGowan: The kind of issue I’m talking about can istration. maybe be illustrated by an example well removed At Scripps you will find many scientists who from CalCOFI and the oceans and us here. It’s been serve on all sorts of government advisory panels. said, and I believe it’s true, that there are ten million Their advice is sought. I can’t guarantee that their homeless farmers in southern Brazil. They represent advice is often listened to. But at least there are rec- a very serious problem to the Brazilian govern- ognized avenues. We have a political process - as ment-a social problem and a potential political some people say, we have the best Congress money problem. The policy of the Brazilian government can buy. has been to cut down Amazonia to provide farms Nevertheless, it is important that tensions exist in for homesteaders. Whether the farms will work or our society, partly because of the environmental not, I don’t know. movement, which does a spectacular job in raising Most scientists, of course, are horrified by the these issues and bringing them forward and pushing idea of destroying Amazonia: look at all those spe- on them. You must continue. There are other sides cies; look at all those beautiful trees; look at the who are also pushing forward. Somehow, in the ten- diminishment of diversity. And after all, the jungle sion, we work out a political process that is, unfor- provides oxygen . . . and on and on and on. But tunately or fortunately, the best one we have. what about those ten million peasants? That’s the Scientists do have a voice. There are many ways kind of advice that I’m very dubious about. How do to get our views known, both to the federal govern- you make a judgment about what it is we want to ment and to the state government. Because we are a preserve? Which do you choose - those poor bas- public instution, all of our information is paid for by tards who are trying to live and survive, or a bunch the state of California or the federal government, of parrots? (laughter) and is available to everybody. Sulnick: I would like to make a response to that, And it is up to you, the environmental movement, although I’ve never represented parrots before. to interpret it one way; it is up to other people to (move laughter) interpret it another way; it is up to the federal gov- I think that that is not a good argument. ernment to interpret it a third way, etc. I agree with Clearly - and Boyce’s remarks, I thought, were per- John McGowan: we scientists do not have the nec- fect on this - the problem is overpopulation. We essary right to say that this particular scientific result rarely talk about it because it’s verboten - then you has that particular effect on society. There is no rea- get birth control, and that’s a big deal. It’s hard son why we should be any wiser than anybody else enough to raise money to run a 50l(c)(3), much less in that regard. I think we have to have a certain tell your audience they’ve got to deal with birth con- amount of humility about what our results mean. trol. But clearly that’s the problem. And clearly, the We should make them uniformly available. We

51 SYMPOSIUM: GLOBAL CHANGE AND THE MARINE ENVIRONMENT CalCOFl Rep., Vol. 31,1990 should take our teaching responsibilities seriously it, which is the side of the status quo. Again, this is and get those messages out to as many people as not necessarily bad, but without question has its possible. It just does not work in any other fashion consequences. that I can see at the moment. Let me ask Boyce a question: You clearly said that Thorne Miller: I want to say that where scientists scientists need to become involved in the environ- may represent only one viewpoint, it is an important mental dialogue. How do we environmentalists one. And if you back off from the decisions, then the meet them halfway and bring them in? Because decisions will be made with an unbalanced view- many times I hear from scientists, “You environ- point, because other people will not be afraid to mentalists really don’t want to hear from science, come forth and offer solutions. and you reject our information if you don’t agree So I think scientists do need to take that extra step with it. ” to suggest solutions and make sure that the infor- Thorne Miller: We do want to hear from scientists. mation that they have provided is used. And the And it’s not just I as a scientist who wants to hear solutions finally chosen may not be the ones that the what scientists have to say. The Oceanic Society has scientists suggest, but these must be considered as a service called a technical assistance program, and part of the equation. we hear from small groups around the country who Sulnick: I want to ask you, Boyce, a question, but say, “We need technical information. We’re working first I’d like to say to Professor Frieman: the infor- in a vacuum. We feel that this particular coastal issue mation is available, but it’s really not accessible, and is an important one, but we don’t have the expertise it’s a mistake to believe that it is accessible. And I to fight this battle. ” want to explain why that is. A good example is the proposed Monterey Bay When I took my job at American Oceans, my Marine Sanctuary off the coast of northern Califor- salary was cut more than half- dramatically more nia. A group came to us and said, “We want the than half of what I made as a lawyer. And that’s sanctuary to be as large as possible. In fact, we think neither good nor bad. I just want to point out that it should go up to the Farallons, but the Marine environmentalists don’t have a lot of money. And Sanctuaries Office [of NOAA] is resisting that. We when you don’t have a lot of money, you don’t have don’t have the scientific information. ” We contacted a lot of support. So I don’t have an associate, as I had some scientists at Santa Cruz and found that, in fact, in the law practice, and two secretaries, and a whole they believed the same thing that the environmental support system, to whom I can say, “Would you group did. So we can facilitate that interaction, get analyze this for me, please. I need it by tomorrow,” those people together, and get both the environmen- and then have it. talists and the scientists talking to the decision Instead I would have to spend a lot of time on my makers. own- which is not a bad thing: I certainly am ca- Sulnick: Would anybody who is a scientist like to pable of doing it - gathering that information and respond to that? making it accessible, first to me and then to my Frieman: Just one quick point. Jackie Parker is the audience. I don’t have that process built in, because head of our Public Affairs Office. She will respond I can’t pay for it. And I can’t do it myself and still to phone calls from anybody, and will try as hard as run the organization and do what else I have to do in she can to put people from the public in touch with life. So while it is clearly available, it’s important for Scripps scientists. If you want information in terms you to understand that it’s not really accessible. And of reports, she will facilitate getting them. That’s until it becomes so, it will only be used by those one of the reasons we have this office. I admit that it who can pay to access it, which is of course industry. has a very small staff, in terms of the issues you’re That’s not bad; it’s not unfair. But that is the way raising. But this is an important point: we are a pub- it is. lic institution. We have a responsibility to get our So what environmentalists always have to do is inforination out. We have such an office to try to work overtime, which is not a problem, but it’s still help, and when anybody calls Jackie, to the extent very hard to compete with the opposition that will that she can (she works terribly hard and has an spend millions of dollars in accessing the informa- overworked staff, as we all do), she will endeavor to tion and then interpreting it. And in order to rebut get the information to you. This resource is available the opposition’s interpretation, you need to have an to you. expert witness or you really lose credibility. I just Sulnick: Professor Scheiber, your remarks about want to make it clear that although the information the watershed events and the turning points were may be available, if it is not easily accessible, it is wonderful, I thought. My question is, do you see going to be used by the side that can afford to access one coining up in the near future over the global

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crisis that is perceived among us, and is the role of is burning down around us, so to speak, meaning science in that to help initiate it, or simply to respond that the problems are very serious on the global to what comes out of the grass roots? level, is it not possible that the role for an institution Scheiber: I agree with Dr. Frieman’s remarks en- like Scripps in the twenty-first century should be tirely. I think we’re not in a good position now, different than it was in the twentieth and the nine- either as scientists or as people who study science in teenth, and that it should be dedicating itself to set- the policy process, to advance this enterprise. ting this agenda rather than just doing objective There’s not agreement on what has to be done. research? There isn’t the kind of agenda that existed in these Frieman: I believe that my role here is to try to put turning point situations: one reason that they were Scripps on a course that I call Scripps 2000. I and my turning points is that there was an agenda. Scientists colleagues, as you can see by the color of our hair were able to guide policy in a given direction, faced and the lines in our faces, are ready to depart this with a specific problem. system. There are a huge number of young people In the case of global change, there is enormous out there who have to be trained to take over posi- variation in how the dangers are perceived. Ob- tions of leadership. viously, from the discussion in this room, there is I regard the agenda of global change as really the enormous variation in the degree of confidence with future of Scripps in the next decade and beyond the which scientists from the various specialities or year 2000. I’ve made that clear to our faculty. We are those seeking an ecological view approach the prob- hiring new faculty, and we’ve organized new re- lem. Contrary to the moments in time that I men- search divisions along those lines, and so on. That’s tioned in the post-1945 history of fisheries our internal business, but nevertheless it is part of a oceanography, the issue isn’t really being presented major agenda. As a leading institution in the United to scientists in a coherent way today. They are being States, we must take a scientific leadership role. asked to give it coherence. So as I say, I share Dr. I then ask the other question: What else can we do Frieman’s rather gloomy assessment of how well as scientists? Suppose that this ecological disaster equipped we are at this moment to deal with it. I really does creep up on us. I think that we are in the think we’re floundering, not in this room alone, but very early stages of understanding mitigation. We in the professions. understand global mitigation - reduce fossil fuel Sher: Does that lead you, then, to the conclusion use, switch to alternate fuels, switch to nuclear, that the government should not be taking any steps? switch to solar, switch to fusion when it comes Or if that’s not your conclusion, what steps should along; eliminate chlorofluorocarbons; we’re doing government be taking, given this diversity of opin- that. We believe that methane is a major greenhouse ion about the problem? gas; we have no idea what to do to control methane. Scheiber: One possibility would be to accept that, Methane might be a much larger piece of this whole remote as we are from a coherent conception on program. We can reforest. As 1 go further on in this which a great majority can agree, continuing sup- list we find ideas that are more speculative: dispose port of the still-fragmented approaches is war- of CO, by burying it in the oceans; ferry huge ranted. I’m not preaching standing back and doing amounts of sulfur dioxide into the atmosphere to nothing at all. reflect sunlight; put satellites up like venetian blinds Sher: Do you think there should be more money for to control the sunlight. We as human beings have research, or should there be some. . . . enormous intellectual power, and we haven’t Scheiber: Yes. More money for research on lines thought through the mitigation problem at all. We that are directed toward a better and more coherent have not really encouraged ourselves or let ourselves definition of the problem. do that. I think we have to get on with thinking Sher: But how about the argument that the problem about mitigation quickly, because it’s a neglected is overtaking us and that if we wait for the results of part of the agenda, and it may be one way to cope the research and don’t take the preliminary steps it with the problem if it comes on us rapidly. will be too late to deal with the problem? Scheiber: I’d like to ask Assemblyman Sher, just to Scheiber: Yes, I agree with that. You’ll be happy to clarify his question to me: What would you put a know I support you on the forests; I think there moratorium on in this area? It’s clear what you do should be a moratorium and not just a study. about cutting ancient trees, but would you put a Sher: I wish you were a member of the legislature. moratorium on all economic activity? Sulnick: My last question is for Professor Frieman. Sher: That’s the kind of proposal, obviously, that I If one assumes - which you may not, so part of my resist, and it’s not going to be acted on. But what question is to ask you to clarify this - that the house would the scientific community do in terms of

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trying to promote a decrease in chlorofluorocar- “Look, I need your staff to do this,” and some do bons? What kind ofdrastic actions? Is that important and some don’t. How do you handle it? enough? A number of bills were introduced this Thorne Miller: I try to call NOAA. year. There was a perception that this is a very per- Sulnick: As I said, in my mind this is a matter of sistent problem, affecting greenhouse gas as well as economics, because all of those data are available, causing a problem with the ozone layer. Is science but access is clearly not, from a practical point of ready to mobilize and say in a unified voice that this view. Obviously the information is public; anybody is something we must do now? We know it’s impor- can go and look at it, but one needs time to go and tant. We don’t know how serious the global change look at it, to write an analysis ofit. problem is, but we know this will help, and there Scheiber: Actually, some of the federal data bases are other reasons to do it. have been privatized; they are not public. A lot of That’s what we’re looking for: we need help when the data that researchers want have to be purchased we take these initiatives - there are people in the today. It’s a very serious problem. legislature who respond to this problem, who’ve Sher: Let me tell you how we do it in government. been contacted about it, who read about it, and so We use moles. . . . There are a lot of agencies, and they drop bills into the legislative hopper. That’s not necessarily for scientific information, particu- what you elect us and pay our minimal salaries to do larly when the executive branch is in the hands of (laughtev) -introduce legislation. So even though one political party and you’re in the other party, or we don’t know whether the problem is really going just generally in another branch of government. to overwhelm us, there are good reasons to take There are individuals in the various departments and some steps anyway, and this is one of the strategies. agencies who know a lot of things, but they are I want to see that kind of thing put together with the under constraints. And yet they have friends on the help of the scientific community. outside, and they feel strongly about an issue. It’s So often in the legislature, as the moderator was not just whistle blowing, but it’s a lot of other saying, when proposals are made, well-financed in- things. They make knowledge available, and then dustry and business groups who will be impacted the politicians on the other side can start digging by the proposals use science in the other way. They without revealing their sources. find scientists who will say, “We don’t know the I suppose there are people in the scientific com- answers, or we’re not sure of this. It’s not good sci- munity who are under similar constraints, but who ence you’re doing here. ” So in the legislation we set can furnish what they think is revealing information up scientific advisory panels. That’s why there are a that ought to be known and acted on. lot of them in federal government; they are not put Sulnick: The second part of this question is rather there in order to collect the information to lead to interesting, the way I’m reading it. If a member of action. A lot of them were put there to prevent ac- the faculty at Scripps makes a public statement, is tion. And that’s what’s a little frustrating for me. that attributable to the institution, and how does the Sulnick: We have received a lot of written questions institution respond to that? from the audience, some of them general and some Frieman: As I said, as far as I’m concerned, any addressed tO individual panel members. I’ll present member of the Scripps faculty can make any state- the general questions first. ment in public that he or she would like. That is The first one reads: “Much scientific information their right as citizens of this country. They cannot, and opinion is housed in institutions with political however, make the statement in the name of the in- constraints, i. e., agencies and other governmental stitution, because there are 1,200 people here, and bodies. How can that expertise found in individuals maybe 1,199 disagree with them. So the individual be released into the public arena in the same way that simply should not speak for the institution. That’s Scripps allows its scientists to speak as individuals, all we’re saying. while taking no position as an organization?” McGowan: I can confirm what Ed has been saying. This is directed to anybody on the panel, but first In my thirty years here at Scripps Institution, I’ve I’d like to comment. I read this as a two-part ques- never felt any constraints against getting up and tion. One part has to do with the myriad data that speaking my mind in public. (laughtev) As a conse- exist in different agencies like EPA or NOAA, data quence, I’m not very often invited to do so. But it is that are really astounding to try to tackle. I’ve tried, a serious matter. One should not claim special con- and I’m sure Boyce has tried. The way I handle it is sensus expertise or give the impression that one is that I call up my local congressperson and say, speaking for the institution in general.

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Thorne Miller: I see a little more to the question. I Part of what I tried to say earlier is that none of think it also is asking about the highly competent these folks will argue that it’s not important to live individuals in government agencies who perhaps within the bounds of conservation, whether you de- cannot speak out. I think that this is not so much a scribe it as the allowable biological catch, which is a question of cannot. I think Jim Hansen, for instance, term now in the fishery management area, or as the showed us that they can speak out; they can express maximum sustainable yield (which, by the way, is a an opinion that is not necessarily representative of term I abhor, but I’ll argue that some other time). the agency, in his case NASA. But it’s complicated. But nevertheless, the pressures are strong. When they give testimony in Congress, very often Scientists, don’t you start taking a conservative that testimony is reviewed by members of the view of what these numbers are because you’re con- administration who say, “No, you can’t say this; you cerned about other things. If you really think that have to say that. ” I think that the individuals in the the system can support a 1,200,000-t0n removal of agencies who feel strongly about this need to speak pollock for the next year, that’s the number you out and say, “Give us our chance.” should give to the policymakers. And you ought to Sulnick: The author of the next question begins by do everything you can to support it and to stand stating the function of this meeting - to talk about behind it. Other people, for whatever reasons, will what society needs from science in preparation for try to increase or decrease that. What we want from global change-and then asks, “Ought not the you folks is the ground truth. We want it stated very question be rephrased to the following: Because hu- succinctly, very clearly. And we want you to stand mans must exploit the environment to some extent behind it. And then the political system will start to to survive, how can science help society to continue work. exploitation responsibly in the face of global Other than that, I don’t know how to answer this change?” question. There are groups of bioeconomists and A follow-up question is addressed to Bert Lar- there are social scientists coming into the realm. So kins: “Since the Alaskan factory tanker fleet will far, at least in my experience, the mix of biologists, need to overfish the resource in the next two years economists, and sociologists has not really worked to make mortgage payments, what do you feel is the very well. There are social and economic aspects to first step needed to implement effective manage- almost all of the actions that my members and any- ment to allow maximum sustained yield of the body else fishing for a living have to face up to. resource?” We’re starting to hear such things now as, our fleet, I think the general flavor of these questions is because of its bigness, is responsible for increasing good, because we are a society based upon the con- the suicide, abortion, and divorce rates of Kodiak. sumption and exploitation of resources. Some of our That’s pretty powerful in a political arena. And how resources are nonrenewable, and some take a long do you defend against it? Maybe it’s right; I don’t time to renew. But all of them, once used, show a know. consequence in the global scheme of things. So the Sulnick: Isn’t it true that this debate about exploi- question becomes for all ofus, in one way or another, tation is really directed at each one of us? Because all How do we begin to to retrofit our thinking so that of us are living a lifestyle dependent upon exploita- we can sustain maximum yield? tion. We all have mortgages; mine isn’t $30 million, Larkins: The members of my association are think- but I have to make the payment, and I assume every- ing about this themselves. They are sort of the last body else has an equivalent. So isn’t this the ques- of the great pioneers. They’ve generally been op- tion: How do we no longer exploit and still manage posed to limited entry; the ocean is the last place to carry on with the work of living? This is not a that’s a common property resource; we can all go question directed just to fisheries. It’s a question di- out and compete- “The cream floats to the sur- rected to all of us. How do we change our lifestyles face,” we hear daily. So government, stay out ofour so that we no longer dump oil into the gutter that hair; just give us the biological maximum. If the runs into the coastal zone, or so that we conserve quota’s a million tons, okay, that’s all we want. Some miles per gallon? It’s a generic question. of us will make it, some of us won’t. Larkins: I’d like to make one more comment about That’s a bit simplistic; some of the viewpoints are this. For a long time, even when I was working in starting to change, particularly among those with Seattle in the sixties, 1 had some interface with the $30 million mortgages. They are starting to wish folks down here in the tuna industry. The supersein- they had bought a 100,000-ton share five years ago. ers were just coming on line; overcapitalization was

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being talked about. But people always knew there Sulnick: I want to tell you. We were walking along was another opportunity. They weren’t sure what it on the same path, step for step, and then you went was, but “Okay, if we’ve overcapitalized this one that way and I went this way. I said, “That’s right, ” and we can’t make it in the tuna industry, sardines and I assumed your next sentence was going to be, will come back, mackerel will come back. ” We, and “We’re going to have to change the way we live.” even folks on the East Coast, and in the tuna fleets, To me that’s obvious. looked to the Bering Sea. There was a 2 million ton- McGowan: I don’t want to. (laughter;) It took me plus potential resource up there that was already too long to get here. marketable if you knew how to market in Asia, be- Sulnick: That’s the way it goes. To my mind, you cause there were Asian fleets up there exploiting it. I don’t cut down the rain forest to maintain your life- know of factory trawlers that were built in New style. You can’t kill all the dolphins to maintain your England when things were rather depressed there lifestyle. You’re not going to have drift nets to main- (they have since gotten worse), and some people tain the lifestyle. To me, that is insane. Nothing said, “Well, we may end up in the Bering Sea.” That personal. (laiightev) was their out. I think that this is the real debate. And we, each There isn’t any out any more. The whole com- of us, will obviously determine the outcome. Be- mercial fishing industry - at least the groundfish, cause to the degree that you say you’re not involved, coldwater fishery - has suddenly, for the first time, you’re teasing yourself- to put it mildly. met itself coming around the corner. And we are just Sher: Conservation is not going to come about vol- now starting to face up to this. The political-scien- untarily, by individuals. It’s going to be done by tific system that’s been working within this com- regulation. The best example of that is what’s hap- munity for thirty or forty years has suddenly come pening in the South Coast Air Quality Management to the same conclusion. And I think the shock value District. We’ve been reading about the new air basin is such that we really haven’t sorted it all out collec- plan and the 120 or 130 regulations that have just tively or independently. There is no place else to go. come out in order to comply with ambient air qual- We have tuna boats in the North Pacific fleet. We ity standards. have some of the converted 200-foot seiners: they These regulations - such things as not being able took the seine table off and put trawl winches on. to barbeque in your back yard, and what kinds of They don’t do very well, but they are probably paints you can apply to your structures - are going doing better than if they’d stayed in the tuna indus- to affect everyone’s life. A tremendous range of ac- try. Many of the vessels that entered the Alaskan tivities will be controlled. There will be very serious groundfish fishery in the last ten years were designed attempts to cut down the number of vehicle miles to do something else when they were built. King traveled, because all of the gains we’ve made in crab boats had stern ramps installed and have be- cleaning up the internal combustion engine - and come draggers. I suppose there are rattails and sau- we’ve made a lot of gain on points of basic science - ries and perhaps one or two other species out in the have been wiped out by the increased number of middle of the ocean, but even FA0 now says that vehicles on the road and the number of miles that there are no great untapped resources left. they are traveling. So there are going to be very McGowan: I’d like to address a couple of points, stringent restrictions on how you can use your per- one that you just raised. And that is, How do we sonal automobile in the future. This will affect our continue in the style to which we are accustomed lifestyle, and it will affect all ofus. But it won’t come and not exploit nature? I think this has been treated about voluntarily. Ifwe had to rely on its being done many times before. One of my favorite philoso- voluntarily we wouldn’t make any gains. phers, Eric Hoffer, wrote an essay about that sub- Scheiber: Following along this line, the Bering Sea ject-a very good one. I think the answer is, we is an interesting case in which there are also conflict- can’t. We can’t do it. We’re going to have to exploit ing uses, and terrible controversies that arise. Not nature in order to live well. only is it the place of last resort in the dreams of It’s the same question, or a very closely related fishermen who are depleting stocks elsewhere, it has question, as the one about the ten million Brazilian also been an arena for enormous international ten- peasants. Amazonia has to be exploited in order to sion and a huge investment of American prestige provide those people with the minimum adequate over the years - first to keep the Japanese out, then life. That at least is the argument. I don’t know what a leading motivation for the 200-mile zone. There is we do about that. a vast history there, and virtually overnight we

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might see it all go to a point of extreme danger with of us who live there. And we know it, even if not oil rigs and tankers out there (a statement that intellectually. doesn‘t sound as paranoid as it might have sounded So I don’t think the issue of global change is a fad. before Valdez; we all understand that a little better But in this question, the word fad is being used to now). mean politically popular. It’s politically viable for So that’s the kind of controversy in which - in the moment. The problem doesn’t go away, but its answer to you again, Byron - choices are clear. We attractiveness goes away. And then do we stop pay- can put off the day of involuntary gas rationing for ing attention to it? That’s the question. And how do another week by opening up all those wells and pro- we ensure that that doesn’t happen? Even if it’s not ducing a week’s supply for the nation out of the in a politician’s best interest to promote x, but x Bering Sea. That’s a very discrete kind of situation. needs to be promoted, how do we continue to pro- The global change problem is not that kind of a mote it, especially when we’re talking about global discrete situation in which you can bring either sci- change? entific or other kinds of expertise to bear as intelli- Sher: I think there are a lot of fads that politicians gently and as rapidly. So that’s where the perplexity respond to, one of which, for example, is whether is. we ought to amend the Constitution to make the Thorne Miller: I followed along your path, Bob, burning of the American flag illegal. There are cer- and made that same divergence with John Mc- tain powerful emotions, and people read the polls. Gowan. But I also want to remark on the comment But in the environmental area, I don’t believe that’s about voluntary change. I think that there can be the case. I think that where there are serious prob- some impact from voluntary changes. A good ex- lems and there are people working on them seri- ample of this is the recycling issue. That is some- ously, we don’t get that kind of political mileage out thing that we have had to push policymakers hard of working on them. There are too many organized on, and we’ve done it by people voluntarily showing forces against us. . . . that they can recycle. The local government refuses That may not be entirely true; you’re going to to legislate recycling requirements, saying residents find an environmental initiative on the ballot next won’t go along with them. So the citizens take the year. One of the gubernatorial candidates is sup- matter into their own hands and establish voluntary porting it, and it deals with three major areas. One recycling programs. The programs prove success- is offshore oil activity; a second is pesticides; and a ful, so the local government decides, “Okay, then third relates to various aspects of the global warm- we’ll put in recycling regulations. ” ing problem. It’s true, people are using that because Sulnick: I’ve got four more questions that are pretty they detect a public awareness and involvement, and good, and I’d like to get one or two of them in. The they think it will benefit them politically. But those first one is to you, Assemblyman Sher. issues are not fad issues; they are all serious. And “Environmental fads come and go, yet global we’ll talk about them for a while, and the initiative change develops over a long time - decades, years. may or may not pass, and if it passes it may not be What mechanism can be implemented to ensure implemented very well to deal with the problems. It long-term commitment of federal and state agencies isn’t that the politicians will walk away from it be- to document and understand environmental cause it’s no longer fashionable. change?” That same idea was raised in the New Yovk McGowan: I’d like to make a comment in my role Timestwodaysago. . . . as a scientific answer man. I think there is absolutely Sher: Let me see if I understand: is that money for no question among reputable scientists that concen- research again? (laughtev) tration of CO, in the atmosphere has been increas- Sulnick: You’ve got it. Next question. ing. We know that for sure, as well as we know Sher: I’m for it. I always support money for anything. And so it is for other gases that affect research. radiative transfer of heat and energy from the sun. It Sulnick: This is an interesting question to me be- is a virtual certainty that the increase in these gases cause I don’t personally believe this is going to turn is going to cause a warming of the global atmo- out to be a fad. When one gets sick, there are symp- sphere. Almost every reputable scientist I know of toms, and the earth is showing a lot of symptoms. If believes the theoretical physics behind that argu- you don’t see them, come to L.A. for a day and ment. breathe the air. That habitat is so stressed out that it What is uncertain is whether or not we have de- really is not supporting the quality of life for those tected that warming as yet, because there are, after

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all, cycles in climate, and this recent warming may be, we should put the brakes on as quickly as possi- be just part of a larger cycle. ble. I don’t think that means that we should drive What is even more uncertain, and in my opinion, our cars every other day; I think that means we virtually unpredictable now, are the consequences of should drive cars that don’t add to global warming, that warming. We don’t know what the magnitude which, my information tells me, are technologically is going to be; we don’t know where it’s going to feasible. But not yet politically acceptable. But that take place; and we don’t know how environmental debate would just go on and on. So, using the pre- systems, especially biological systems, are going to rogative ofthe chair. . . . respond. That’s where the real questions are. And Question from the floor: When we get down to that’s where the uncertainty is. But it is a phenome- the specifics, what are they really going to be, and non - there’s no question - that’s going to happen. are we prepared to advocate moving to nuclear It’s not a fad. power? Sulnick: In your view, is it a phenomenon we Sulnick: As moderator, I don’t know that that should actively seek to stop? ought to be what we are debating, because everyone McGowan: I wrote something down here earlier: Is has a point of view. As a quick response, I think our present knowledge sufficient for major political there are alternatives that would promote an energy action, or are there uncertainties enough to delay policy based on conservation and renewable fuels, action? Isn’t that the question? and would not jeopardize the nation with the poten- Sulnick: Precisely. tial devastation of nuclear energy. McGowan: Yes and no. There is sufficient knowl- If I can avoid having a debate on this I would like edge right now to say that we absolutely must do a to, simply because it’s four o’clock and I have one better job of understanding what the consequences more question I would like to present to the group of change might be. We must tune up and become before I thank you for your participation. much more elegant and sophisticated in studying The question is: “How much money is the United change. Change as compared to what? What kind of States government giving CalCOFI to investigate a baseline do we have? We must know the magnitude the amount of contaminants the United States sends and direction of change, and it must be compared to to Mexican territorial seas through the California what the ordinary state of the system is, on a global Current?” basis. We really have very little of that information. Mullin: I should point out that this question was And yes, we need action in terms of, if you’ll excuse translated from Spanish. Obviously there is a legit- me, more research. imate concern on the part of our Mexican colleagues Sher: But you wouldn’t take people’s cars away that there is a flow of water from the coast of Cali- from them, based on what you know? fornia to the coast of Mexico. I don’t know that it McGowan: Not yet. will be possible for the panel to come up with a Sher: You wouldn’t make them travel fewer miles number, but it’s certainly worth thinking about. in order to cut down the carbon dioxide yet? McGowan: The other part of that question is how McGowan: Oh, sure. much does the federal government support Cal- Sher: You .would support a law that says you can COFI studies relevant to this? There is no support only drive your car every other day? from the National Science Foundation or the Envi- McGowan: But there is already a lot of carbon diox- ronmental Protection Agency; only NOAA sup- ide in the atmosphere. ports the program. Most of the support comes from Sher: What I have to respond to is proposals that are the state of California. made to at least try to stabilize how much CO, is Sulnick: I have one final statement: I would like to going into the air. And what I want to know is thank the audience. You’ve been really attentive, and whether you are going to support me. obviously involved, and that made it a much better McGowan: The economic consequences of some of discussion than it otherwise could have been. the suggestions might be very severe. The cure Mullin: In closing, I suppose it’s obvious to point might kill the patient, and that’s what I worry about. out that a very large number of people have contrib- Sulnick: Let me add one other point of view. It uted, both to the symposium today and to the seems to me that while we study things we should CalCOFI meeting for the last three days. I’d partic- do so from a safe, or relatively safe, perspective. ularly like to thank Mary Olivarria, Sadie Gonzalez, And if the environmental changes that are taking Lari Maczko, Debbie High, and Kitty Haak, who place are as potentially destructive as they appear to have probably walked a hundred miles between here

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and our office to try to get all of the details straight- in their lives in the last two weeks to keep them ened out. occupied for quite a while. We very much appreciate Again I’d like to say that I’m grateful to our panel, their coming here. particularly those from the Bay Area and Sacra- And finally, again, thank you to the audience for mento, who have probably had enough disruption your participation and support.

59

Part 111 SCIENT1FIC CONTRl BUT1ONS

SCHEIBER: CALCOFI‘S EARLY YEARS ColCOFl Rep., Vol. 31,1990

CALIFORNIA MARINE RESEARCH AND THE FOUNDING OF MODERN FISHERIES OCEANOGRAPHY: CALCOFI’S EARLY YEARS, 1947-1964

HARRY N SCHEIBER School of Law (Boalt Hall) University of California Berkeley, California 94720

INTRODUCTION tinuing to make important contributions to an ad- “Throughout the 1950s and into the 1960s,” as vancing oceanography even after four decades of McHugh (1970) asserted in a study surveying trends corporate existence-in the face of all the odds as- and accomplishments of U. S. fishery research, there sociated with the modal “life cycle” of such scien- arose tific and other academic enterprises (Knauss 1990). a growing realization that the simple prewar concepts of This historical perspective on CalCOFI in its scientific fishery management are not very useful in prac- early years (1947 to 1964) will focus on two impor- tice and that successful fishery management must be tant aspects. The first concerns how the scope and based on scientific understanding of the resource as it design of CalCOFI research on the California Cur- interacts with all the physical and biological variables in rent, and on the Pacific Ocean more generally, were its environment. originally formulated - that is, how the marine sci- The shift in concept of which McHugh wrote was entists and fisheries management specialists, indus- truly profound, not only in its impact upon the sci- try leadership, and state and federal policy officials entific approach to fisheries management but also in defined their research strategies and future needs in its transformation of the ocean sciences. What oc- 1947-49. The state of American ocean research curred during the decade and a half following World when the project was first designed will also be dis- War I1 was nothing short of a methodological revo- cussed. The second aspect concerns the dramatic de- lution. research was reunified with velopment of the range and modes of scientific work in physical and chemical oceanography and inquiry in the early years of CalCOFI research. The meteorology, and a new holistic approach to the focus will be especially upon how a great conceptual study of ocean environments emerged; researchers divide in ocean science was perceived and then dra- sought to analyze the processes of change in com- matically breached, opening the way for modern plex biotic communities rather than to study seg- ecosystemic studies of the oceans. mented processes or small geographic units of the deep seas (Scheiber 1986; McEvoy 1986). DEFINITIONS AT THE FOUNDING: A The coordinated marine fisheries and oceano- COMPREHENSIVE DESIGN FOR “THE PACIFIC graphic studies that would become known as Cal- RESEARCH FRONTIER” COFI (California Cooperative Oceanic Fisheries The enterprise that became CalCOFI was set in Investigations) played a crucially important role in motion in 1947, when, as the successful culmination this transformation of marine studies, participating of efforts by a small group of scientists, government centrally in the great methodological advances. The officials, and industry leaders, the California legis- contributions associated with CalCOFI research lature approved a special tax on commercial sardine cover virtually the whole spectrum of techniques landings (McEvoy and Scheiber 1984). On the in- and subject areas of research in what has become dustry’s initiative, the tax revenues were designated known as fisheries oceanography in the post-1945 specifically for research on the causes of the sardine era, from improved trawls and a new approach to decline - commonly also termed the sardine deple- comprehensive egg and larval studies in the late for- tion - which was then troubling the state’s impor- ties to the modern-day applications of remote sens- tant sardine fishing industry and the processing ing. It is a remarkable thing, moreover, that despite plants and canneries that it supplied. some rough spots along the way, CalCOFI has Landings of pilchard (California sardine) by the evolved successfully and survived in full vigor, con- state’s fleet had slumped from their phenomenal levels of the 1930s, when the sar- Permission to reprint is granted to U.S. government agencies All rights other- dine fishery in the California Current was said to be wise are reserved by the author and CalCOFI. [Manuscript recewed February 27,1990 ] one of the world’s most intensively exploited marine

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Figure 1. Photograph taken on March 14,1947, at Stanford University at a meeting of representatives from the California Department of Fish and Game (I),the California Academy of Sciences (2),the South Pacific Investigationsof the U.S. Fish and Wildlife Service (3),and the Scripps institution of Oceanography (4). Front row: Milner B. Schaefer (3),John L. Kask (Z), Frances N. Clark (l),John F. Janssen (l), Julius B. Phillips (l),Osgood R. Smith (3),and Donald H. Fry (1).Back row: Harald U. Sverdrup (4),Oscar E. Sette (3),Wilbert M. Chapman (2),Carl L. Hubbs (4),Robert C. Miller (2), Elbert H. Alhstrom (3),Richard S. Croker (l),and Kenneth M. Moser (3).

fisheries, and had then fallen disastrously in only a As a political achievement, the background story few years following the war. Thus the sardine har- of CalCOFI is one of intrigue and rare skill in the vest dropped from the peak of nearly 800,000 tons arts of persuasion, coalition engineering, and insider in the 1936-37 season to only half that level in 1945- political trading in response to alarm about the sar- 46, then fell again to only 130,000 tons in 1947-48 dine crisis. A small cabal of industry leaders, SI0 (Radovich 1981). and state fisheries laboratory scientists, USFWS sci- The special tax funds were turned over to a joint entists, and political leaders in the legislature put industry-science committee, called the Marine Re- together the sardine research program idea in a se- search Committee. It was this group which, in the ries of meetings in the winter of 1946-47 (figure 1). course of administering the sardine research funds, The key players, at first, were Wilbert (Wib) M. would some years later formally establish the Chapman, curator of fishes at the California Acad- CalCOFI enterprise as the coordinating body for the emy of Sciences; Montgomery Phister of the Van research it was helping to sponsor for the U.S. Fish packing corporation; Carl Hubbs of SIO; and, soon and Wildlife Service (USFWS), the Scripps Institu- coming onto the scene in a major way, Harald Sver- tion of Oceanography (SIO), the California Fish & drup as director of SIO; Richard Croker and Frances Game Commission, and, on a much smaller scale, Clark of the state Fish & Game Commission scien- Stanford University and the California Academy of tific stafc and Roger Revelle, then in naval service in Sciences (McEvoy and Scheiber 1984). Washington but soon to return to SI0 as associate

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director.’ [Numbered notes begin on page 79.1 It tvansfovmation $the scope and content ofscientijic method was an elaborate political dance. They first lined up in ocean science - that is, the basic concepts of marine the reluctant support of the notoriously individual- studies, and not merely a dramatic expansion of the istic fishing boat owners and corporate executives in geographic scope of studies in the eastern Pacific. the sardine canning firms - men who were, as Phis- One instrumentality ofthis vision was to be struc- ter called them, “captains and individualists, ” never tural, invoking the coordination of agencies and the prone to join ranks with one another or anyone else, collaboration of multiple disciplines. The sum of the suspicious of the ivy-towered scientists and the re- enterprise (funded by the MRC and the cooperating source managers, such as Clark, who spoke the lan- agencies) would be made far greater than its parts guage of regulation and restraint.2 by coordinating the skills, ships, equipment, and The organizing committee - which one insider knowledge of the state marine fisheries laboratory, late termed “the proto-MRC,” because it was pred- SIO, Stanford University’s marine laboratory (the ecessor to the Marine Research Committee (MRC), Hopkins Marine Station), the California Academy the body established by the California legislature to of Sciences, and the federal agency. Beyond that, the administer the new program - forged an uneasy al- sardine project would share data and plan its work liance between Scripps Institution’s scientists, who jointly with the ocean scientists based in the fishery were committed to pure research in physical and agencies of the other West Coast states and British chemical oceanography and in marine biology, and Columbia.’ Not least important, the sardine project the fisheries scientists in the state and federal agen- could complement - and in fact from the outset it cies who were interested in “mere” applied manage- was coordinated closely with - the oceanographic ment concept^.^ The leadership also somehow and fisheries work being started in 1948 under Oscar overcame much of the long-standing mutual mis- Elton Sette’s leadership, in a Hawaii-based federal trust between the state scientists and the federal tropical tuna project.H agency. (“The Federals,” Chapman wrote early in the course of politicking for the project, “are ac- The State of Marine Reseauch’in the Pacific to 1947 tually wondering whether or not they wish to get There has been vast growth in the last forty years, any further involved. . . .”4) since the California cooperative project on the sar- Not least of the unlikely achievements of a win- dine began, in knowledge of the Pacific Ocean in all ter’s whirlwind lobbying and alliance-building, the its aspects -marine biology, ocean chemistry, geo- committee obtained not only industry’s consent to a physics, and meteorology. It is astonishing to con- special tax on sardine landings, earmarked for the sider how little, by contrast, was known in 1947 of research project, but also the legislature’s agreement what California marine scientists at that time liked to pump large new appropriations into the SI0 to call the “Pacific Ocean fisheries frontier” (Pacific budget to support the work, especially to operate Fisherman 1947). The sardine crisis was only one seg- three new large-scale research vessels. These ships ment, albeit a dramatic one with enormous eco- were donated to the University of California, for nomic impact, of a vast congeries of interrelated use by SIO, by act of Congress and cooperation of mysteries about the Pacific. the U. S. Navy. But they were also the special gift of The archival records and some scattered scientific Comdr. Revelle, who from his post in the Bureau of publications of the era reveal that to a remarkable Ships managed to get the service’s approval to trans- extent the small community of West Coast ocean fer these newly decommissioned warships to the scientists had a keen understanding that this larger university, as well as congressional appropriations and more comprehensive web of unsolved mysteries for their reoutfitting for research.’ had to be attacked if ever the resources for adequate It was the sardine crisis that set all this under way, research came to hand. They understood, in other but from the outset what Chapman and some of the words, how little was known about this ocean sys- others had in mind was a much more comprehensive tem and the precise nature of its dynamics. The pat- push for what he termed “high seas research on a terns of the currents, the basic bathythermography, scale far beyond anything that the United States has the ocean floor in the deep-sea areas, meteorological undertaken or thought about in the past.”” Chap- phenomena in relation to biological systems and hy- man’s language referred not only to research drography - all these were scarcely known, despite throughout the Pacific basin: for underlying all the the brilliant formulating of what we may term the early deliberations of the MRC scientists, as I will “right questions” by pioneering figures such as seek to show here, was a vision that foresaw the Sverdrup and his associates at SIO, William E

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Thompson at the University of Washington in Se- salmon was most important in providing the direc- attle, Clark and other California state scientists in tion and intellectual framework of Pacific studies, fisheries work (mainly on the sardine), Albert Herre the fisheries management scientists resorted to an on tropical fisheries in the western and South Pa- emphasis on harvest theory and the concept of max- cific, Sette of the USFWS research office in Califor- imum sustainable yield, indicated by harvest vol- nia, and a few other notables.’ ume (output) in relation to inputs (“fishing effort”) What of fisheries research more narrowly? A star- (Russell 1942; McHugh 1970). This almost exclusive tling limitation was that West Coast research had emphasis, responsive to the needs of the fishery in- long been confined largely to the inshore areas. Even dustries and becoming the basis for some successful the most basic questions remained unanswered for management programs (most notably, the halibut some of the great pelagic and anadromous species. effort undertaken in 1931 by Canada and the United No one knew, for example, how many distinctive States, with Thompson in charge), meant a loss of populations of tuna inhabited the Pacific, where momentum for the more problematic and difficult they spawned, or, even at the grossest level, how work of dealing with ecosystemic relationships. abundant they were. There was uncertainty even Marine scientists did not lose the vision of ecosys- about whether the warmer Pacific waters and in- tem study, to be sure; fishery experts trained under shore areas of the western and South Pacific region Thompson himself, for example, later recalled read- had stocks enough to support fisheries on a sustained ing in their journal groups at Seattle the studies by commercial basis.l0 Similarly, in the North Pacific, Hjort and other pioneers in the ecological style. But apparently no one, at least in Canada and the United during the interwar years in the United States and States, had the slightest idea whether or where Asian elsewhere, the requisite money, gear, technology, and North American salmon intermingled in the instrumentation, ships, and personnel were entirely high seas (the Japanese did have some fragmentary lacking for work in this mode (Herrington 1988; data that they kept secret), or knew what events in Scheiber 1988). the high seas most affected the stock during the life Given greater resources in scientific personnel and cycle (Herrington 1989; Scheiber 1989).l1 funding, even within the existing limitations of re- Identifying the relationships between the condi- search technology, much more could have been tion of fishery populations and their ocean environ- learned, but research on environmental relationships ments (including such aspects as nutrients, food to fisheries remained fragmented, small in scale, chains, chemical properties of host waters, currents lacking in spatial scope or intensity. Pacific Ocean and weather, patterns of predation and interspecific studies on the West Coast of this country were, in predation, etc.) had been in the minds of fishery sum, almost unbelievably impoverished. The bril- scientists since well before the end of the nineteenth liant but scattered achievements of an era that century. When the first of the great Scots coastal stretched from the Wilkes Expedition in the early fisheries surveys was established in the 1880s, for nineteenth century to the Albatross and Cavnegie example, even before the Challenger reports were voyages of 1900 to 1931 had been followed by a dec- published, the stated objective was to understand ade in which only one American-flag research vessel the relative impacts of human activity and environ- (the E. W Scvipps) was dedicated to basic oceano- mental conditions on fisheries (Deacon 1990). The graphic research in the Pacific. Only a handful of importance of such ecologically framed study had scientists did offshore research, and many of that also been recognized in the coastal and seabed fish- small number were in agencies whose funding was eries research in Scandinavia and northern Germany based solely on their mission of conducting applied at the turn of the century, best exemplified in the research on coastal fisheries management. The lan- work ofJohan Hjort. Without question the environ- guishing of this American research effort, because ment’s relationship to commercial fishing and its im- the resources were not there, compounded the very pact on marine resources had motivated the real difficulties associated with the state of available formation ofICES at that time (Idyll 1969; Dymond technology for deepwater study (Shor 1978; Schei- 1948). But research on these lines had generally been ber 1986). frustrated by the limitations of technology, gear, and In retrospect, however, it seems evident that at the funding: the oceans were too vast and impenetrable. important West Coast centers of study - SI0 for the As a result, in the interwar years, 1918-39, the chemical and physical sciences, and secondarily for focus of commercial fisheries research had shifted biology; USFWS and the California state agency for radically. Led by William Thompson, whose theo- commercial fisheries research; the University of retical and applied work on sardine, halibut, and Washington for salmon and halibut research and

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oceanography; and Berkeley and Stanford for zool- Rather, there was a shared awareness of what needed ogy and biology - the small cadre of ocean scien- to be done to get the work started in the Pacific.” tists, numbering perhaps thirty at most, understood The best of the fisheries management scientists in with remarkable insight what were the most impor- 1947-48 were already keenly aware of what they tant gaps in empirical knowledge and methodology. needed to learn in areas where they had little or no Precisely for this reason, as we will see, they were data - on problems such as interspecies competi- able to reach broad agreement as to what an agenda tion, or the relationship of nutriment levels to juve- for expanded study ought to look like, and which nile survival rates, or the role of upwelling, which kinds of inquiry would be likely to yield the most had been explored from a meteorological perspec- knowledge of ramifying ecosystemic relations.” tive in the brilliant early Pacific studies of Sverdrup In this context, the decision to launch the Califor- and his associates at SI0.14 Once the prospect of new nia sardine project constituted a remarkable depar- funding, gear, and ships was at hand, the scientists ture in the history of Pacific Ocean research - a quickly produced their wish lists. landmark in the reestablishment of a major Ameri- Perhaps this is in itself unremarkable; all good can research presence in Pacific science. It put funds professionals have some kind of wish lists ready at in the hands of Pacific marine scientists at levels that hand, in the happy event that funds should suddenly were ten and more times the revenues for research become available. The historian will find, however, that they had enjoyed in the previous two decades, much more than random or disparate priority lists and it made possible the inauguration of deepwater in the archival records of the CalCOFI project and research by several new ships, well equipped with of its progenitor the Marine Research Committee, the latest gear, that extended by an enormous mag- or in the personal correspondence of scientists such nitude the capabilities for ocean research. At about as those who masterminded the California push for the same time as the California legislature author- research funding: Chapman, of the California Acad- ized formation of the MRC and the cooperative sar- emy of Sciences, who principally orchestrated the dine project, the U.S. Congress moved to correct political moves, put his intellectual imprint on the what had become a scandalously embarrassing de- research proposals, and, rather miraculously, re- ficiency in support for Pacific oceanography by es- cruited the fisheries industry to the cause; Carl tablishing the Hawaii-based tuna research project Hubbs, Sverdrup, and Revelle of SIO; Sette of the (POFI), the scientific work of which was initially federal agency, joined in 1948 by John Marr, who under the direction of Sette, with Schaefer in charge would succeed him in charge of the U.S. Fish & of biological studies. Wildlife Service South Pacific Fishery Investiga- Although the crisis that galvanized California was tions; and Frances Clark, that remarkable woman the sardine’s critical decline, the national motivating who, by her studies over many years in the Califor- force was a larger geopolitical concern expressed in nia state agency, had established herself in the front congressional debates: the concept of Pax Ameri- ranks of and was a pioneering ad- cana and more specifically the intermeshing ambi- vocate of stronger management constraints. ’’ Many tions of the U. S. Navy, the Pacific fishing fleets, and of these scientists were also associated to varying the fish-canning industry to establish the American degrees with the overlapping effort to obtain presence in Pacific deep-sea waters before other na- congressional action to establish the POFI project in tions, friendly or otherwise, had recovered enough Hawaii. The published sources and surviving per- from the devastation of war to stake out claims that sonal correspondence that express scientific thinking would preempt U. S. interests (Scheiber 1990a). in the West Coast community of fisheries specialists reveal important common themes and a core of com- Understanding “One Ocean As a Whole”: The mon objectives. Pacijic Vision There were also some important cleavages, to be A striking feature of the California effort in this sure, within the scientific community - the diver- surge of new activity in Pacific research is the fact gent interests of the biologists versus the physical that the community of West Coast ocean scien- and chemical oceanographers, and a very clear de- tists -however fragmented in other respects - had marcation between applied and pure scientists. their agendas fairly ready in hand when the political There were also important differences of view moment for action arrived. This is not to say that among the fisheries-management scientists as to there was a “Pacific Oceanographer’s Manifesto,” how heavily to rely upon landings data for evaluat- or the equivalent of some priorities handbook, to ing the condition of the stocks. Considerable per- which any and all ocean scientists might subscribe. plexity was also evident as to how, ifit could be done SCHEIBER: CALCOFI’S EARLY YEARS CalCOFl Rep., Vol. 31,1990

at all, to build on the insights of early-day fisheries move far off the California coast and to “make the ecologists - led by Johan Hjort and followed up by entire Pacific our oyster.”” Michael Graham and others who had sought to re- Looking back on this element of “original in- late environmental conditions to fishery dynam- tent,” as one may term the vision that animated ics-as a way of getting beyond Thompson’s CalCOFI and other Pacific projects in the late 1940s, harvest-yield approach that was so dominant at the we can see how it became a permanent part of the time (McHugh 1970; McEvoy and Scheiber 1984). program design for the next four decades. This But a key element of shared understanding, evi- widely shared understanding that the Pacific Ocean dent in the various agenda ideas that were sent back required study in its entire scope - that fishery and forth among the Pacific Coast scientists and that problems could with great profit be intensively ended up as working policy documents for the di- studied in relatively small regions, but that the nat- rection of new projects, was the sense that the scope ural variables affecting abundance and condition of ofvesearch ultimately must be the Pat+ Ocean and not such regions might be located only through study merely discrete geographic regions and segments in of vast areas - has given impetus to the elaborate which one species or another dwelt. This was a vi- coordination of far-flung projects, both American sion that went beyond solving even a crisis so omi- and international, that produced the vastly more nous and disturbing as the sardine decline that was complete empirical portrait of the Pacific Ocean sys- then occurring. tem that has been achieved in the last forty years (see In October 1946, for example, scientists from the Miles et al. 1982). various state fisheries management agencies of the West Coast, together with a representative of the federal agency, had formally proposed research “to The Ecological Vision establish the relationship between oceanographic The more timeless element of the new vision as- fluctuations and the concomitant fishery phenomena sociated with the sardine project’s design related, . . . [requiring] a continuous record of conditions in however, to the fundamental conception of the both fields: physical oceanography and fisheries. ”lh ocean science enterprise: it was an ecological vision, The U.S. Navy Hydrographic Office quickly en- and it departed radically from the prevailing mode dorsed this view of a need for “extensive synoptic of twentieth-century ocean fisheries research, and oceanographic information about the waters off the indeed ocean science generally, especially in Amer- Pacific coast of North America, ” and especially “ex- ica. It was a return, in effect, to the older tradition panded investigations of the departures from normal of studies exemplified by Hjort and others who had oceanic circulation” - an endorsement that well re- sought to integrate fisheries management and ma- flects the direction of thinking that prevailed among rine biology with broadly conceived environmental West Coast oceanographers at that time.” For this research. was precisely the view that Chapman, the SI0 Again, the archival records reveal a scientific vi- group, the federal scientists (Sette, Marr, Walford, sion set forth with remarkable clarity and presci- and Ahlstrom), Robert Miller of the California ence. An exemplary document in these records, Academy of Sciences, and others expressed con- though by no means the only one that might be stantly during the hectic period of planning for the singled out for citation, is a statement of the research sardine studies in 1947-48, preliminary to the form- design first prepared by Roger Revelle in late 1947. ing of CalCOFI. He contended for a new conceptual framework of This large strategy for research was recognized biological study in relation to ecosystems - “to eloquently by one of oceanography’s leading fig- make dynamic analyses . . . of the processes in the ures, Columbus Iselin of Woods Hole, in a con- sea, that is, the cause and effect relationships which ference address at SI0 in 1951. Appraising the affect sardine production. . . . ”’” importance of the California group’s sardine studies “In the past,” Revelle continued,” and other new research, Iselin commended the West Coast scientists for giving substance and hope to the oceanographic research has been concerned primarily idea that it might be possible “to understand at least with the description of aver.age conditions prevailing in one Ocean as a whole.”’’ Similarly, Roger Revelk the sea. The investigation upon which we are about to and others at SI0 often voiced the view after 1948 embark poses a new and more difficult problem, that is, that their new capabilities-reflected in the ships, of studying the nature and causes of variations from the gear, funds, and technology that were then at their average conditions.” The present is a good time to start disposal - permitted their institution’s scientists to such an investigation, because obviously we are in a pe-

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riod of major departure from the average conditions, at That message was not lost on the sardine industry least insofar as the distribution of the sardine population cosponsors of CalCOFI, who became painfully is concerned. aware that “their” problem was becoming part of an In attacking a problem of such magnitude all possible ever-ramifying scientific enterprise that was coming scientific tools and methods will have to be employed. It to focus upon ecological systems. The fishing and will be necessary first to describe as completely as possi- cannery interests had to ble the existing oceanographic and biological situations; be reassured, on many oc- second to establish empirical statistical correlations be- casions, that basic research on a broad conceptual tween the various environmental and biological factors; basis would eventually produce important practical and third and most important, to make dynamic analyses results.” where possible of the processes in the sea, that is, the It was vital that the sardine project succeed, Chap- cause and effect relationships which affect sardine pro- man declared, because it had been “log-rolled duction. Wherever such a dynamical analysis of a partic- through by a small group of far-sighted men in the ular aspect of the problem can be made, a great saving in industry [who were] far ahead of the main body of time required for a solution will be effected over the the sardine industry in their thinking,” and if the “brute force” method of statistical correlation which re- work succeeded in producing results, the whole in- quires a long series of observations for validity. . . . dustry would fall into line; if it failed, the “die- The sardines cannot be treated as isolated organisms hards” would prevail and “our work is very apt to living in a vacuum. The investigation must be an inte- grated one in which proper weight is given not only to be set back for a generation. ”” the currents and other aspects ofthe physical environment The subversive content of the new vision was not but also to the entire organic assemblage including the lost, either, on the applied fisheries management sci- plants and animals which form the food chain of the sar- entists -especially Frances Clark, who at times ex- dines, their competitors for the food supply, and the pred- pressed deep frustration with the way that analysis ators, including man. . . . of complex systems could divert attention from the intense commercial fishing effort that she believed The vision that Revelle set forth entailed, in sum, to be the real culprit of the sardine-depletion piece, interdisciplinary research in a holistic mode: its fo- whatever the other variables and their subsidiary ef- cus was to be the ecosystem. As had already been fects (McEvoy and Scheiber 1984; McEvoy 1986). learned from experience in the earth sciences, he The subversive side of the ecological vision not- wrote, “far more productive results were obtained withstanding, its constructive side would have a by complete analysis of all the factors which exist in profound influence upon the direction - and the a particular situation than by a statistical treatment most renowned achievements - of ocean science in of a few factors in many situations.”’3 Similarly, the ensuing decades. Beyond that, however, this Sette of the USFWS had written that to study the holistic or ecosystemic approach that was encapsu- sardine dynamics properly in relation to ecosys- lated by Revelle in 1947 and eloquently endorsed and temic change, it would be necessary “to set up a later imaginatively pursued by others - Chapman, program on a basis that will cover much more of the Schaefer, Marr, Clark, Walford, Ahlstrom, Mur- sea area along the Pacific Coast [than had previously phy, and other intellectual leaders of the CalCOFI been studied] and will run through enough years to enterprise and related Pacific studies of that era- establish the average conditions and discover what was the precursor of the more comprehensive move- effect the deviations from the average condition have ment in modern science toward ecosysternic re- on the recruitment and availability. ’v4 search designs. The major shift toward such holistic Thus Revelle’s presentation in 1947 set forth a pre- analysis of systems would occur only in the 1960s, cise and unambiguous agenda for research in a “par- amid the new political concern for “environmental- ticular situation, ” the California Current. (And in ism,” when it was also reflected in the reconcep- this respect it described exactly the mode of research tualization of the public policy approach to that would actually be pursued by MRC and Cal- environmental monitoring, regulation, and risk as- COFI for four decades.) But the ecological vision sessment (see Fleming 1971). that he expressed also had a “Subversive” side, as Fully fifteen years earlier than that, however, in good science and interpretive theory in other fields the late forties, the ecosystemic concept and the re- of study usually do: this subversiveness was to be search designs it inspired became one of the truly found in the implication that the ecosystem, and not glittering achievements of the CalCOFI program - merely the sardine dynamics as one part of that sys- essential to its foundations from the outset, and tem, was the truly interesting and enduringly im- manifest thereafter in the pursuit of California Cur- portant subject of inquiry. rent studies.

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CALCOFI’S EARLY PROJECTS AND THE lation to availability [also to California F&G]. 6. EMERGENCE OF A NEW FISHERIES Dynamics of the sardine population and fishery [a shared OCEANOGRAPHY research area, for all participating agen~ies].~’ CalCOFI celebrated its fortieth anniversary in 1989, but strictly speaking the project dates from the The scope of the plans, and also the way in which formation in 1948 of the Marine Research Commit- they reflected a comprehensive view of the sardine tee (MRC), under terms of the legislation of the population’s dynamics, indicated a substantial in- previous year. The actual research supervised by the crease in personnel and gear as well as ships. In the MRC was set in motion by the Technical Commit- months that followed, John Marr of the federal tee (composed of four scientists charged by MRC agency coordinated planning with SI0 and the state to oversee the work at sea) in the early weeks of fishing management scientists to deploy the new 1948.27The research was inaugurated in February SI0 ships, a refitted federal vessel (Bluck Douglas), with a hastily organized voyage into waters south and the N. B. Scojield (also, later, another California from the SI0pier, to make some quick visual obser- state vessel, Yellowjin, a refitted naval ship). The plan vations of sardine movements and (it was hoped) that emerged from the talks called for observation pick up a few samples in the offshore waters from stations across a grid that went 400 miles off the San Diego south to Punta Abreojos. This mission coast, with probes for collecting nutrients and other was conducted by the E. Pi! Scvipps, the heroic little materials at depths of nearly 3,000 feet. Expendi- wooden research schooner that had been the main tures by MRC, from the sardine tax revenues, reliance of the Scripps Institution scientists in their included $50,000 to the U.S. agency for ocean- upwelling and other high-seas studies in the 1930~.~* ographic work, $25,000 for its egg and larvae stud- The portrait of ocean phenomena that we can now ies, and additional sums for gear and personnel for obtain from space satellites and the advanced gear of the other agencies.3’ modern oceanographic vessels reminds us how far The SI0 leadership sought out additional labora- the study of the oceans has moved, conceptually and tory personnel to process samples as they came in technologically, in forty years. That little ship beat- from the research vessels, and new oceanographic ing down the coast, in its 1948 quest to locate some gear (bathythermographs, barographs, plankton sardine runs, was - in its conception, in its gear and nets, flowmeters, high-speed collectors, sonar de- instrumentation, and in the limits of what it might vices, etc.) was purchased with the funds from the hope to accomplish - much more akin in many re- sardine tax.3’ There soon emerged a lively competi- spects to the exploration and science associated with tion for trained personnel among POFI in Hawaii, Captain Cook’s voyages than to the oceanographic the new MRC projects, and other Pacific research studies of our own day. centers (especially the salmon research center at Se- The California Fish and Game research ship N.B. attle); SI0 began training people for specific posi- Scofield followed soon after, with an April voyage in tions available on several of the newly expanded quest of evidence of sardine stocks off the Baja Cal- projects (Scheiber 1986). ifornia coast. Meanwhile the SI0 scientists and navy Thus was set in place the extraordinary station personnel worked at a frantic pace to modify and plan of the sardine research program under MRC. outfit the two former war vessels that had been The station plan was the heart of the continuing turned over to the University of California for SIO’s CalCOFI studies and their extended longitudinal research at sea.”9 data series, covering an area of some 670,000 square A formal agenda was set out at the April 1948 miles (figure 2). The program provided for detailed meeting of the MRC, at which Dr. Robert Miller, sampling and testing of ocean water to determine chairman of the Technical Committee, presented a hydrographic conditions. Samples were collected to six-part program that included the following lines analyze chemical and physical properties of the of research: waters; the volume and composition of nutriments; and larvae, juveniles, and adult fish (Ahlstrom 1950). 1. Physical-chemical conditions in the sea [assigned to The oceanographic sampling program and also the SIO]. 2. Organic productivity ofthe sea and its utilization [also to SIO]. 3. Spawning, survival, and recruitment of larval and egg sampling cruises were initially con- sardines [assigned to the federal Bureau of Commercial ducted at 122 stations Over the vast Fisheries]. 4. Availability of the stock to the fishermen nia Current area; they continued on this (behavior of the fish as it affects the catch) -abundance, basis for Over a decade, until they were reduced to a distribution, migration, behavior [assigned to the Cali- pattern of quarterly cruises, albeit over a more ex- fornia Fish & Game Division]. 5. Fishing methods in re- tended ocean area (Wooster 1949; Murphy 1960).

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------STATION PLAN OF THE COOP E RAT 1V E SARDINE RESEARCH PROGRAM 0 0

SHOWING POSITIONS OF STATIONS 0 OCCUPIED DURING SURVEY CRUISES FEBRUARY THROUGH SEPTEMBER.1950 45

40

35

30 70. 235

90 261 13: 60 90280 13070 25' 13: (10'

135" 130' 125' 1200 115'

Figure 2. The CalCOFl station plan used for cruises in 1950. The numbering system was planned so that the station lines were 120 miles apart, and individual stations were 40 miles apart. Extra stations were added in regions of particular interest for sardine work. (See inside back cover for basic plan.)

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The extent of startling changes in scope, com- ready for sea.”35This latter view prevailed, so in late plexity, and scale in California marine research February, Cvest, Hovizon, and the state vessel N.B. doubtless helped to prompt the June 1948 reflections Scojeld began the work, with the N. B. Scojeld of Carl Eckart, who succeeded Sverdrup as SI0 di- newly equipped with hydrographic and plankton rector that year: “The individual scientist, working collection gear, bathythermographs, echo sounding in seclusion,” Eckart declared, “is apparently a thing equipment, and other instrumentation from SIO, as of the past.” Although he was uncertain that this well as high-speed collectors, standard plankton was “going to be good for science,” Eckhart nets, and flowmeters from the federal agency.36 thought it was necessary (if it were to work at all) It is worth noting that within the year the state that the new research projects “be led by people who sardine tax funds administered by MRC were also have a comprehension of the past. ”” being used for some gear and operating expenses of Given the deep and continuing involvement, and the USFWS cruises that were augmenting the Cali- leadership, as the sardine project ramified, of distin- fornia project with surveys off the coasts of Oregon guished fisheries scholars and oceanographers who and Washington -perhaps the first such instance in had already excelled in highly individual research, it American governmental history of significant appears in retrospect that the success of this experi- grant-in-aid money flowing against the established ment in Big Science-mode organization and coordi- currents, i.e., from the state to the national govern- nation was built on exactly the critical foundation ment, rather than from Washington to the ~tate.~’ that Eckart prescribed. There is no gainsaying, The sardine project initiated by the MRC, accord- however, that “seclusion” as a way of scientific life ing to a Los Angeles Times account based on a pub- had been discarded. Indeed, nearly every great ac- licity release from SI0 in June 1949, had quickly complishment in MRC-CalCOFI research in this become the “biggest fish hunt in history. ” Revelle era reflected the intricate collaborations of agencies termed the project, in the same report, a “fine- and institutions, the crossing of disciplinary lines, tooth combing of coastal waters, ” asserting that it and the cooperative relationships that developed be- indicated that “the State has decided that it is almost tween the scientists at sea and the laboratories to as important to develop and conserve our sea food which their findings were sent for analysis. It was, resources as it is to develop agriculture. ” Sympto- quintessentially, what Eckart termed the new-style matic of the larger agenda that by then was explicitly “organized scientific effort. ” emerging, however, was Revelle’s further observa- tion concerning the longer-run objectives that could An Expanded Design and a New Name be realized through the new project: “The outcome With the cooperative deployment in 1948 of ships of the all-out sardine research, ” he declared, “is vital and scientific personnel, financed by a mingling of to thousands, but OUY ultimate aim is to obtain scientijic agency and University of California funds, the Ma- data without which we can’t hope to assuve a maximum rine Research Committee’s sardine project was fully sustained yield of food fvom the ocean” (Los Angeles under way. The project moved forward, however, Times, 1949, italics added). under the dark clouds of continuing crisis in the By mid-1949 the larger contours of the project sardine industry, as the decline in catch continued. were fully etched: they included the more compre- Indeed, the state marine fisheries scientists, while hensive dimensions of the research in the realm of lending the full weight of their efforts to the new pure science, subsuming and in some respects be- research, were at the same time pushing hard for ginning to overshadow the applied fisheries man- authority to strictly limit commercial fishing for agement concern. The state agency, whose scientists sardine and to close several ocean areas.34 throughout the entire early history of the MRC ef- Aware of the political controversies about man- fort were embroiled in controversies over whether agement decisions underlying the sardine project’s to halt sardine fishing as a way of stemming the start-up, Revelle (then returned from the navy and precipitous decline of the resource, expanded their serving as associate director of SIO) opposed delay- traditional agenda of studies that were based on ing the grid cruises for any reason. When Marr, the standard harvest-yield and input data.38 Croker, USFWS representative, suggested that more time Clark, and the agency scientific staff had long pur- was needed to outfit the ships they had assigned to sued recruitment research, including work on dis- cruise the northern part of the grid, Revelle replied tribution and methods of conducting census surveys that “financial and political reasons” alike made it of the nursery grounds, and they had continued to “almost essential that we should start the first measure abundance by using bait-fishery statistics. cruises with our own [SIO] ships as soon as they are Aided by the MRC initiative, the agency had also

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mapped out additional exploratory cruises on the Coast of North America we will learn much about fishing grounds, designed to produce data on “cor- the regime of ocean currents and temperatures. ” relation of physical and biological oceanographic Other major issues that were suggested by the stud- conditions with sardine distribution” and to im- ies completed to that date, Revelle stated, were prove methods of locating the stocks, studying The use of zooplankton as indicators of water masses school habits and other behavior, and studying “re- and diffusion. lations to other species (mixing of schools).”3” The development of methods of collecting post-larval In the case of the federal agency - reflecting the stages of a variety of pelagic fishes. approach to larvae and egg research pioneered by Studies of the role of oceanic birds as pelagic fish Ahlstrom and Sette before the war and then contin- predators. ued under Sette, Ahlstrom, Walford, and Marr dur- The unexpectedly large populations of many species of ing the late 1940s -the initial cruises in the northern pelagic fish other than sardines in off-shore sub-surface range for the MRC sardine project demonstrated waters. even more immediately how the scientists’ agenda The discovery that electrical signals can be sent up or went well beyond the narrow issues of sardine man- down uninsulated hydrographic cable . . . [raising] the agement. Thus the 1949 progress report of ths fed- possibility of almost revolutionary developments of methods for continuously measuring sub-surface tem- eral scientists on the sardine project (which the peratures and other variables from equipment towed be- agency subsumed under the title Expanded Pilchard neath the surface at normal cruising speeds4’ Research Program, within the framework of its South Pacific Fishery Investigations then headquar- Over the course of the period ending in mid-1950, tered at Stanford University) highlighted the issue the broad categories of research that would be pur- of possible interspecific competition on the sardine sued over nearly a decade had become well estab- grounds. Analysis of the plankton and egg samples, lished. As summarized in a report by Ahlstrom the agency reported, indicated that anchovy larvae (1950), the SI0vessels gachered data in several major were distributed in roughly the same areas as sar- areas of investigation. The first was physical ocean- dine, and that an abundance of hake and jack mack- ography, including studies of upwelling, transport erel had also been found in areas where their of water, chemistry of the water, and “the causal presence had not previously been re~ognized.~”Pre- mechanisms behind the circulation the ocean.” Sec- dation on the sardines was suggested as a factor lim- ond was the study of phytoplankton, concerned iting their population, and in any event, the report with evaluating the “crop” of marine plants and par- continued, “the other fishes may be competitors for ticularly “the relation of fluctuations in the produc- food and space” with the sardine. If any single theme tivity of marine plants to physical and chemical pro- was hammered home, it was that of the ramifying cesses in the ocean. . . and the effect . . . on the scope of the research inquiry: populations.” Third was the study of zoo- plankton, especially its effect on survival rates of In addition, valuable data are being gathered on fishes larval and adult sardines. Fourth were marine verte- which are of great importance in the economy of the sea, brate studies, conducted in close collaboration with although not of direct commercial importance. It is be- the U.S. Fish and Wildlife Service scientists, espe- coming increasingly self-evident that the biological, cially in pursuit of what seemed a promising and chemical, and physical studies being carried out with im- dramatic possible breakthrough in understanding mediate reference to the sardine problem will be of tre- mendous value to the study of many other fishes.“’ the dynamics of relations between sardine stocks and other species, especially the anchovy and possi- Similarly, in forwarding a report by SI0 on its bly the saury. As Frances Clark of the state agency cooperative role in the MRC project during the 1949 rather dolefully observed at about the time of this cruises, Revelle scarcely mentioned the ~ardine.~’ summary report: “Scripps is doing the new and Again, he emphasized instead the ramifications of spectacular and appears to get a lot of praise and the early research findings for basic oceanography glory. ”44 (See also Pan-American Fisherman 1950). and marine biology, and not the pressing applied By contrast, it was the state Fish & Game Division (and highly politicized) issue of sardine fishery reg- scientists who were tied down to what Clark termed ulation. The findings in the past year’s work, Revelle “the routine drudgery without much glory, ” work declared, indicated the desirability of exploring the in which “no one is interested . . . and it is without hypothesis that “to a large extent the ocean is the publicity value.”45The state vessels and scientists slave of the wind and that if we gain an understand- continued to pursue research on the lines that had ing of the dynamics of the atmosphere off the West been pioneered by their agency since the 1930s,

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studying distribution, harvest statistics, and sam- nicely into the bureaucratic niche existing in his ples that reflected survival of year classes. Despite agency in the form of its Pilchard Research Program the lack of publicity that irritated Clark, the agency budget category. Responding for the state agency, was also contributing importantly to the grid station Frances Clark suggested “Cooperative Marine Re- program and the building data base of synoptic search Program,” which had the advantage that it oceanographic and biological data (Ahlstrom 1950). “does leave the way open for tying the work in with The South Pacific Fishery Investigations scientists other fisheries” (though she added, “This may or continued, through cruises and in their own and SI0 may not be an advantage”). Revelle carried the day, laboratories, to explore the environmental relation- suggesting on behalf of SI0 that he would “favor ships manifested in the research on sardine recruit- something a little more comprehensive” than the ment and survival that had been highlighted in their title Marr had put forward; hence he suggested “Co- earlier report. Other, smaller, elements of the sar- operative California Fisheries Research Program. ”47 dine research program under MRC included a Soon afterward, the name that was to become small-scale bench project at the California Academy permanent and universally referred to by the acro- of Sciences that involved experiments with sardine nym CalCOFI began to appear on the project’s schooling behavior, and correlation and analysis by publication^.^^ all the cooperating agencies of the commercial catch Throughout the early years of CalCOFI research, statistics that were being generated by government the publicity releases prepared by the University of resource-management agencies in Oregon, Wash- California and other agencies stressed, as did Ahl- ington, and British Columbia, as well as by the strom and Hubbs in reviewing a 1952 public rela- USFWS and the California state agency (Ahlstrom tions statement for radio use, that “although this is et al. 1950). a sardine investigation, the investigation is contrib- This 1950 report indicates how far the orientation uting to a better knowledge of all the fisheries and and guiding vision of the program had gone beyond to a much better understanding of the ocean itself. ”4y the sardine management issue by its emphasis on Similarly, the publicity efforts underlined that the more general phenomena of the oceans. Upwelling ramifying implications for ocean science methodology received full discussion, and there was exten- were also of key significance. Indeed, the radio sive analysis of food supply and food chains, the broadcast release stated that, whatever the fate of relationship of nutrient supply to intraspecific and California’s sardine population and the sardine fish- interspecific competition, and the possibilities of ing industry, “perhaps in the long run, the most mortality associated with disease-producing organ- significant thing about the sardine investigation is isms as well as predation by competing species in that it’s demonstrating the feasibility of large-scale fishery populations. The report declared the emerg- fisheries research. ”50 ing character of the sardine research program to be The rapid development of an ecosystemic ap- proach to research issues and to the actual design of studying the sardine in its environment in order to under- the MRC-CalCOFI program did not entirely dom- stand how this environment - physical, chemical and bi- inate the project’s history in the earliest years. As ological - affects the survival of the sardines when young mentioned already, a major theme- really part of and their distribution (availability) when they are of com- the contextual fabric - was the continuing tension mercial size. , . . We are studying the sardine ‘at home. ’ , , . in the larger arena of state politics, centering on To date, little more than a good beginning has been made on the study of environmental conditions. Yet it is rather whether or not strict management controls - even certain that before we can hope to predict fluctuations in suspension altogether of commercial sardine fish- abundance of the sardine fishery we mustjnt investigate the ing-should be imposed on the industry. McEvoy environment thovoughly enough to undevstand the ejfects ofphys- (1986) has argued that the scientists at SI0 and ical and biological yvocesses OM the savdinepopulation.4“ USFWS in effect willingly ran interference for the industry, which was heavily aligned against the The ethos that by then pervaded the leadership’s cause of regulation. An extension and continually conceptions of the broad direction and ramifying ramifying expansion of the research project, he con- significance of the research was expressed in corre- tends, was entirely congenial to scientists who con- spondence among the project scientists in 1949 re- sequently enjoyed unprecedented funding for basic garding a name for their program. John Marr, who research; and it worked to frustrate the intentions of had become chief of the South Pacific Fishery Inves- the California Fish & Game Division marine scien- tigations of the federal agency, proposed “Cooper- tists, who firmly believed that overfishing, whether ative Sardine Research Program,” a title that fitted alone or in conjunction with other forces, was the SCHEIBER: CALCOFI’S EARLY YEARS ColCOFl Rep., Vol. 31,1990

instrumental factor in the sardine’s decline and pos- uninterrupted time series (which, of course, would sible imminent disappearance. be continued for five more years on the original grid It seems to this writer that a somewhat different pattern) on temperature, salinity, currents, and scenario was being played out - that the basic other variants.54 oceanographers and federal agency scientists consis- To this list of accomplishments should have been tently regarded it as their role to generate good data, added the remarkable breakthroughs in geology work out the best possible research design, pursue that came out of a notable exploratory cruise pro- the leads that scientific judgment suggested (how- gram in 1952-54. The SI0 ships cooperated with ever they might ramify the work), and let the po- the Charles H. Gilbert of the POFI project to inves- litical branches of government decide about tigate the waters between Hawaii and the eastern regulation. To allow MRC or CalCOFI to be split Pacific. In addition to locating rich new areas for apart by differences on an explosive policy issue tuna fishing, the cruises made important discoveries would be to sacrifice the harmony and possibly the about the seabed configuration east of Hawaii (Sette survival of a precious and productive scientific en- 1952, 1955). terprise. In other words, the separation of science The last finding that was summarized in the 1957 from tough policy decisions -a luxury the Fish & report - that “information on the identity, location, Game Agency was not afforded - was more a natu- and abundance of the eggs and larvae of many spe- ral concomitant of the type of research CalCOFI was cies, including the anchovy, jack mackerel, Pacific undertaking than a matter of the science fraternity’s mackerel, saury, and hake, [had] been obtained an- cynicism or something even more sinister.” nually since 1950”- proved to be of truly determi- A coordinate theme, as the environmental and native significance for future CalCOFI research. As ecosystemic vision came to dominate CalCOFI de- Ahlstrom wrote (1964) concerning these early years sign, was the building up of a record of concrete of data collection: “It was a fortunate circumstance accomplishment in science - the cumulative body of that the sardine was found to have a wide areal dis- research that within a decade after the MRC found- tribution and an extended spawning season. ” The ing had made the California Current probably the breadth of distribution was discovered virtually most intensively studied marine fishery area in the from the outset of the MRC-CalCOFI cruises, and ~orld.~’ the findings indicated a great extent of range and the length of spawning season. These findings in turn The Research Achievements, a Data Glut, and prompted the investigators “to look at large chunks CalCOFI Reorganization of the California Current system off California and Summarizing what MRC and CalCOFI had Baja California rather continuously” (Ahlstrom achieved up to 1957, John Isaacs of SIO, John Marr 1964). of USFWS, and John Radovich of the California Because the investigations had been carried into Department of Fish and Game categorized the major so vast an area of the deepwater Pacific, and because research accomplishments as follows. First, sardine the nets had brought up massive determinative evi- spawning grounds had been identified over a much dence that the anchovy and sardine populated the wider area of the California and Baja California off- same regions (and evidence also that there were shore region than had previously been re~0gnized.j~ other species, especially hake and mackerel, that Second, annual estimates had been made since 1950 must interact in some ways with the sardine, their of the number of fish spawning in each of the four food supply, and their activities in the larger physical major areas; eggs and larvae had been estimated an- environment) two things followed. First, the re- nually, as had “the abundance, distribution, and age searchers were led more and more deeply into inter- composition of juveniles and adults on the inshore specific dynamics, a direction of study that would nursery grounds. ” Third, the numbers of adult sar- within a few years lead to conclusions on anchovy- dines had been estimated annually since 1952. sardine competition that would dominate CalCOFI Fourth, studies had been made on various aspects of science for a long time. And second, the scientists spawning, mortality, north-south migration pat- and their agencies were inspired to grapple with the terns, and schooling habits of sardine. Fifth, in the mysteries of the more comprehensive systems of studies of nutrients, the project leaders had con- ocean ecology in ways that greatly transcended nar- cluded that the presence of phosphate and other nu- row concerns with the sardine. trients did not appear to be a factor limiting These developments were reflected in the formal phytoplankton in the region. In the traditional areas statements of program objective that the MRC oc- of oceanographic study, the cruises had produced an casionally adopted during the 1950s and early 1960s.

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In the 1950 CalCOFI progress report, the program this once-great fishery had nearly disappeared, the was summarized as one “to seek out the underlying MRC was still declaring formally that explanation principles that govern the Pacific sardine’s behavior, of this “catastrophic decline” must remain a top re- availability, and total abundance” (quoted in Mur- search priority. Whatever the brilliant achievements phy 1960). In 1954 the objectives had been broadened of MRC and CalCOFI research up to that time and “to include the ... mackerel, jack mackerel, and afterward {including the pioneering studies of an- anchovy.” The program objective was stated still chovy-sardine interspecific dynamics that would be more comprehensively in 1957, as determination of published in the early 1960s), the record was made “what controls variations in populations, size and against the background ofunchecked disaster for the availability off the west coast of North America of California sardine resource. sardines and, as their scientific and industrial impor- The second major area of unresolved work that tance requires, of anchovy, jack mackerel, Pacific was identified by MRC scientists in 1957 was in mackerel, herring, squid, and others”j5(See table 1). descriptive oceanographic studies. Many thousands Accurately reflecting the move into ramifying, of days had been spent at sea; the shore laboratories comprehensive collection of ecosystem data across were staffed at levels which, however inadequate for all the ocean science disciplines, CalCOFI adopted the data that was coming in, were unprecedented in an even broader definition of its objectives in 1961 West Coast biological and oceanographic study; and (Murphy 1963): the lack of very significant year-to-year variation in weather and oceanographic conditions during the To acquire knowledge and understanding of the factors entire period 1947-56 had persuaded the CalCOFI governing the abundance, distribution, and variation of the pelagic marine fishes. The oceanographic and biolog- leadership to maintain the intensive level of repeti- ical factors affecting the sardine and its ecological associ- tive studies at the grid stations, in lieu of shifting to ates in the California Current System will be given a more selective sampling approach (Murphy 1963; emphasis. It is the ultimate aim of the investigations to Ahlstrom 1964). Yet the volume of chemical, physi- obtain an understanding sufficient to predict, thus per- cal, and biological sampling was far outstripping the mitting efficient utilization of the species, and perhaps capacity of the shore labs to process the data.” manipulation of the population. By 1957-60 the CalCOFI program therefore was in serious danger of sinking of its own weight. The After nearly a full decade of CalCOFI research, scientific vision that had pushed the project into however, it had become painfully evident that vutni- ramifying aspects of Pacific Ocean-wide meteorol- fication of the research, the expansion of studies into ogy and geology, and that had also generated the comprehensive investigations of the ecosystem, was vast volume of accumulating planktonic, physical, something very different from integuution. One very and chemical samples at the La Jolla laboratories was troubling issue was the continuing uncertainty as to now recognized as militating against effective anal- the causes of the sardine decline; in 1957, even after ysis of the relationships in the marine ecosystem. The best minds on the project were agreed on what must be done: Sette, for example, declared flatly that TABLE 1 the project must pause and shift from collection to Marine Research Committee (CalCOFI) Revenues and analysis, with priority to explaining the sardine col- Expenditures by Agency, 1947-64 lapse. The primary task, Sette declared, must be that 1. Total revenues: $1,738,718 adjusted of “connecting up what has happened in the realm 2. Expenditures, by agency: of physics, chemistry and planktonic life in the sea MRC committee operating expenses ...... $104,439 MRC program coordination...... 11,141 with what has happened to the abundance, distri- Grants to: bution, reproduction and mortality of the sardine. ” U. S. Fish and Wildlife Service ...... 598,430 Donald McKernan, director of the federal Bureau of Scripps Institution of Oceanography...... 75,737 California Division of Fish & Game...... 198,062 Commercial Fisheries, also pressed the MRC to Hopkins Marine Station (Stanford Univ.) ..... 85,838 shift from comprehensive collection of descriptive California Academy of Sciences ...... 125,890 oceanographic data to an emphasis on analysis. Fu- 3. Total expenditures by type of investigation: $1,790,261 adjusted Sardines ...... $9 15,563 ture research should be “intensified,” not ramified, Mackerel ...... 332,971 he argued, and should be “guided toward a study Anchovies ...... 219,310 of the inner workings of the ocean-atmosphere Herring ...... 11,381 ‘engine’ ~~60 Squid ...... ~- .~ Source: Financial Record of Marine Research Committee, document Concern for better focus and emphasis on devel- dated Aug. 7, 1964, in MRC Minutes, SI0 Archives. oping new hypotheses - and effective explanatory

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interpretations of the data - translated, predictably sure of success within five years ofhis appointment. enough, into a call for organizational reform. The By the mid-l960s, CalCOFI research had come to a Isaacs-Marr-Radovich report urged such a course, strong, if highly controversial, focus upon the an- recommending a reduction of the MRC’s continu- chovy-sardine relationship and its implications for ing oversight, with CalCOFI “leadership, direction, fisheries management in the California Current. responsibility and authority” to be placed in a four- Meanwhile, however, both through continuing person committee of representatives from SIO, the MRC financial support and the larger influence of state agency, the USFWS, and MRC. Even more the now-traditional CalCOFI ecosystem research important, however, the proposed MRC “represen- agenda, SI0 and marine fisheries studies generally tative” should be a “broad and practical senior sci- in the Pacific continued to examine the fishery entist” who would actively coordinate all the stocks in relation to the relevant ocean environment. CalCOFI research and serve “as an integrative The legacy of CalCOFI, in this respect, carried over force. ”6‘ into the important studies of fishery dynamics and The idea of having a senior scientist play the key management conducted under Schaefer’s direction role of proactive coordinator - an effectively super- by the Inter-American Tropical Tuna Commission visory role, representing the MRC, but with a in the 1950s and 1960s (Barrett 1980). The legacy also professional commitment above all to the integrity carried over into the era’s larger, truly international of the scientific enterprise-was an old one in web of complementary and coordinated research CalCOFI. Indeed, at the very beginning of the proj- projects on ocean fisheries and environment - proj- ect effort, Chapman had wanted such a scientist- ects that included the NORPAC, EPOC, and Inter- coordinator position to be integral, but the few in- national North Pacific Fisheries Convention studies dustry supporters of the research plan would not (see Miles et al. 1982). support the appointment of a coordinator, fearing A shift back to the more focused applied-manage- they would entirely lose their influence on the direc- ment issue had become evident in CalCOFI discus- tion (and perhaps the content as well) of the re- sions even before Murphy was named coordinator. search.”’ With the crisis that CalCOFI faced from an Industry representatives on the MRC, most notably awesome data backlog, and with the sardine prob- Chapman, had of course long pushed for such an lem still unresolved even as a matter of theory-after- emphasis. But the chief proponent in the working the-depletion, it was decided that a coordinator science group became John Isaacs of SIO, who as must be hired “at whatever cost” to facilitate more early as 1959 authored a “Proposed Program in Fish- effective coordination and move the project forward eries Research” for CalCOFI consideration. In this on the lines Sette, McKernan, and the project scien- document, Isaacs proposed applying knowledge tists themselves now wanted.h3 from data already gathered on the sardine to analyze Thus after much political jockeying and further more universal dynamics, focusing on hake, an- pressure from the senior agency representatives, the chovy, saury, squid, jack mackerel, and Pacific MRC moved in November 1958 to appoint Garth mackerel. The resulting theories should be used for Murphy as the first CalCOFI coordinator. Having what Isaacs termed “sophisticated experiments himself authored important sardine studies under (quite unlike the conventional management)” in- the auspices of CalCOFI, Murphy was equally at- volving interventions through commercial fisheries tuned to the applied management mission of the to reduce target stocks that had preyed on other project and to the larger ecosystemic vision that had species or competed for their food. Such interven- moved the project since its outset. Under his direc- tions would amount to an outright “alteration of the tion CalCOFI budget, administration, and alloca- fish population toward the composition of preferred tion of scientific priorities were put in tighter shape, sport and commercial species.” In this view, the and he apparently enjoyed the confident backing of commercial fishery was a tool to be used for the the key MRC members to whom he (and CalCOFI) elaborate and comprehensive bioengineering of the reported.64 California Current (Isaacs 1959). If the vision reached far ahead of both the data and Maintaining Momentum and Pvovid ing a New the available theory in 1959, it was not long before FOCUS:1958-64 the idea of interventionist management in such a If giving new impetus to solving the sardine mode resurfaced in CalCOFI discussion. This time “mystery” and more effectively integrating the ap- the new coordinator, Murphy, along with Isaacs and proach to ecosystem analysis was the coordinator’s Ahlstrom, took the lead. Ahlstrom’s egg and larval dual mandate, Murphy could point to a large mea- surveys had revealed a dramatic increase in the an-

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chovy population, occurring synchronously with temic research, in some respects continuing the tra- the sardine’s critical decline in the waters they dition of collecting and ramifying data while in shared. The anchovy-sardine ratio in the larval col- other respects pioneering new scientific techniques lections, Ahlstrom reported in 1964, had risen from and developing new theory for the Pacific Ocean 3.9:1 in 1952 to 16:l in 1957, 23.6:l in 1958, and system and marine systems more generally. 46.8:l in 1959. (See also Ahlstrom 1963.) Pointing at In the ensuing period of CalCOFI history, which the obvious policy conclusion - that purposeful re- is beyond the scope of this paper, these multiple lines duction of the anchovy might relieve stress on the of study, and efforts to integrate them into modern sardine - Ahlstrom (1964) made a rather imprecise marine science theory, have constituted one element yet telling suggestion: “Until now,” he wrote, of the MRC-CalCOFI legacy. In the last forty years we have been in the role of observers. We have been interdisciplinary studies, taking the entire marine watching what has been happening in the ocean. Whether system as their ultimate subject, have become the we can successfully be participants, shaping the course of standard in marine research; and the new “fisheries the events, remains to be seen. Certainly the latter has oceanography” has come to dominate the analysis been one of the prime objectives of ocean research. of ocean fauna and their environments. These two developments in scientific method and research con- The full policy implications were left to be spelled out by Murphy and Isaacs (1964), who explicitly stitute the most enduring legacy of CalCOFI’s stated to the MRC what had been left unspoken in founding vision and four decades of California Cur- Ahlstrom’s presentation: that, since the oceanic re- rent science. gime had come to favor the anchovy (in some rela- ACKNOWLEDGMENTS tionship to the selective, intensive fishery for The author is indebted to Berta Schweinberger, sardine), a new regime of unselective “trophic level J.D., and Barbara Leibhardt, M.A., Ph.D., former harvesting rather than selective harvest within a Sea Grant trainees at UC Berkeley, for their assis- trophic level” could serve to redress the situation. tance; and to Arthur McEvoy, associate professor of Research findings suggested, they went on, history, Northwestern University, whose doctoral that the process is reversible, either by a protracted period dissertation the author directed and whose scholar- of years in which the environment clearly favors the sar- ship, both his own and that done in collaboration dines and/or by re-deploying man’s effect on the com- with the author, on California marine science and munity in such a way as to favor the sardine. The policy, is indispensable background to this work and practicality of this depends on more definite knowledge is cited in the references. The author also extends of the exact ways in which the two species interact. warm thanks to Deborah Day and others of the On this foundation a proposal that a managed Scripps Institution of Oceanography Archives, Ja- fishery for anchovy should be initiated was quickly net Ness and others of the University of Washington built and adopted by the MRC. This became the Library Manuscript Division, and Gary Lundell of “great experiment” idea, one that roiled the political the University of Washington Archives for their waters within the MRC despite the CalCOFI scien- generous advice on research problems and their in- tists’ apparent consensus that their data and judg- defatigable assistance in use of archival materials. ments warranted it. The idea also proved Archivists of the National Archives in Washington, controversial in fishery policy circles, even in some and of the California Academy of Sciences, San highly respectable quarters in fishery science, and Francisco, also extended very cordial assistance. certainly in the political arena at Sacramento. The My colleagues William T. Burke, University of sudden rise of the fabled, if short-lived, anchovy Washington; David Caron, John Dwyer, and Roger fishery in Peru cast serious doubt over the economic Hahn, University of California, Berkeley; and feasibility of the proposal, undermining whatever James Sullivan, California Sea Grant College Pro- slender political chances may have remained for it.”5 gram, all are owed thanks for their sustained (and The anchovy harvest proposal provided a short- sustaining) interest and suggestions. John Marr, term focus, causing endless trouble in the MRC and USFWS (ret.) graciously provided a careful critical perhaps exposing CalCOFI to the kind of treacher- reading. Responsibility for any errors is, however, ous political crosscurrents that the project’s scien- entirely my own. tific leaders had long feared would result from This paper is based largely upon research sup- excessively detailed concern with applied manage- ported by grants from the California Sea Grant Col- ment issues. But all the while, the participating lege Program (project no. R/MA-30, grant no. agencies carried on the broader mission of ecosys- NA85AA-D-SG140), with funding from the U. S.

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Department of Commerce (NOAA), and from the March 2, 1953, File 80, ser. 121, USFWS Records, Record Group State of California, to the Program in Ocean Law 22, National Archives, Washington.) The next day, however, he backed off and reported to Kask that in speaking with Don Saxby, and Policy, Center for the Study of Law and Society, a prominent packing industry executive in California, he “got the UC Berkeley. This funding was augmented by the impression that it would be extremely difficult and probably im- California Policy Seminar, University of California, politic for us to withdraw.” (Memorandum, Walford to Kask, March 3, 1953, File 80, ser. 121, USFWS Records, Record Group and Boalt Hall School of Law, UC Berkeley. 22, National Archives, Washington.) 5 See Shor 1978. The University of California, thanks to Revelle’s NOTES ingenuity, had- as President Sproul ironically put it - acquired its 0. E. Sette, director of the USFWS South Pacific Fishery Investi- very own navy overnight. Sproul did say, not without some anger, gations project based at Stanford University, was also important in that he would have preferred to have been consulted in advance developing the project design, but he was much more in the back- about the negotiations. But after a timely visit from a sardine in- ground than the others mentioned in the text, so far as the political dustry delegation, Sproul came around and gave his retroactive effort in California was concerned. Prof Revelle has said that Sver- blessing to the new fleet. Sverdrup-Sproul correspondence, Feb.- drup, a physical scientist, himself “didn’t think much about biol- March 1947, Director’s Files, SI0 Archives. ogy,” but Sverdrup was certainly positive toward the idea of 6. Chapman to Vern Knudsen, Oct. 22, 1947, Chapman Papers, UW, working closely with fisheries scientists (Revelle 1986). Moreover, For a biographical study of Chapman’s long and influential career, Sverdrup had himself engaged in studies of upwelling that bore see Scheiber 1986. directly on the issue of nutriment levels and fish spawning and 7. Chapman to Montgomery Phister, Sept. 19, 1947, Chapman Pa- survival (Sverdrup 1948). He later revisited the basic problems he pers, UW; Francis Clark to Carl Hubbs, April 6, 1948, Subject had explored in the late 1930s and had advanced through his entre- Files: Marine Life Research, SI0 Archives. preneurial role in MRC’s formation in 1946-48 (Sverdrup 1952). 8. This project was the Pacific Oceanic Fishery Investigations (POFI); For detail on the history of the project during 1946-48, and es- it was based in Honolulu upon its establishment in 1947-48, and it pecially its political context, with brief analysis of the long-term focused on tropical tuna resources of the Pacific. There were three achievements, see McEvoy and Scheiber 1984 and Shor 1978. A divisions for research, the biological division (under Schaefer) range of significant policy issues and fisheries science of both the being the most important; the others were technology, with a focus 1930s and the later period, especially the 1960s, are covered well in on preservation and processing, and fishing. The significance of McEvoy 1986. POFI is discussed in Scheiber, in press, b. Sette and Ahlstrom (1948) recount the results of prewar research 9. The state of the art was exemplified in the extraordinary book by involving U. S. Fish and Wildlife Service and University of Califor- Sverdrup et al. (1942) synthesizing research to the time in ocean- nia scientists on the relationship of environmental conditions in the ography, and summarizing, in the course of argument, much of the California Current to sardine spawning. Pacific research Sverdrup and his associates had accomplished in Letter ofM. Phister to Chapman, Dec. 29,1950, Wilbert M. Chap- Pacific waters. See also Scheiber 1986. For an example of thinking man Papers, University of Washington Library (hereafter cited as on agendas, see esp. 0. E. Sette (1943). A classic statement of re- UW). search method in the early years of modern fisheries oceanography The phrase proto-MRC was coined by Garth Murphy, in “Sum- is in W. E Thompson 1919. mation of Calcofi,” manuscript report (presented before the Cali- 10 An early proponent of the theory that tuna were abundant in the fornia Marine Research Committee meeting, Balboa, Calif, April tropical Pacific was Albert Herre, author of influential papers on 11, 1963) in Minutes of the Marine Research Committee, Scripps fisheries of that area (see, e.g., Herre 1940). Herre’s influence on Institution of Oceanography Archives, La Jolla; hereinafter cited as American scientists’ concern to explore tuna resources was a pow- SI0 Archives. On the factionalism of pure scientists versus man- erful one, as testified by Wilbert Chapman, who in 1944 termed agers, I have relied on Revelle 1986. himself something of a “disciple” of Herre on that issue. (Chapman Letter of Chapman to Phister, Sept. 19, 1947, Chapman Papers, to William F. Thompson, Oct. 28, 1944, William F. Thompson uw. Papers, UW Archives, Seattle.) On the institutional background Dr. Frances Clark of the California Fish & Game Division char- and shortcomings of fisheries research specifically within Califor- acterized the history of her agency’s relations with the federal sci- nia, see McEvoy 1986. entists as follows: 11 On Japan’s knowledge of salmon, in contrast to the almost-nil un- derstanding of deepwater movements of North Pacific salmon, see In our relations with Fish and Wildlife we went at it backward. Herrington 1989 and Scheiber 1989. A summary of salmon man- Clashed, fought, and finally cooperated so that in general things agement problems is in Larkin 1970. are now running smoothly but we all have to be continually on 12 This will be developed later in the text. Evidence on point is a our guard to avoid new clashes. Man is jealous by nature and manuscript article by D. Huntsman (1949), in which Huntsman scientists or pseudoscientists are no exception, perhaps among discussed the difficulties encountered over many years of his and the worst. others’ research -especially in research until 1934 on herring, and (Letter of Clark to Carl Hubbs, April 6, 1948, Carl Hubbs Papers, since 1934 on salmon - in achieving a useful set of theories concern- SI0 Archives.) ing the relationships of oceanographic and biological research. He Tensions between the state and federal agencies did not disappear contended that “the factors determining concentration of [marine] after the sardine project’s founding. Indeed, some of the top fish. . . is an oceanographic matter.” Huntsman continued: USFWS leadership believed it was the wisest course for them to keep clear of the project and the political crosscurrents of debate It is really an ecological problem, involving the relations be- over proposals for placing strict limits on the harvest of sardine- tween organisms and their environments, between the ocean and proposals that were being put forward regularly by the Fish & the life therein. Twenty-five years ago I visualized it [the prob- Game Division scientists in public forums. In early 1953, the state lem of why fish concentrate as they do] as the problem of limit- scientists openly conveyed their suspicion that USFWS personnel ing factors, ofthe factors limiting the distribution and abundance were not turning over all of their data. Bristling at the charge of ofmarine organisms. I studied such obvious factors as tempera- “secretiveness” in hoarding data from cruises engaged in the sardine ture, salinity and light. . . . (but) made no particular impression. research, L. A. Walford, chief of the Branch of Fishery Biology in The field of study was still too vast and inchoate for easy com- the federal agency, wrote to USFWS Assistant Director Kask: ‘‘I prehension or for solution of the problem in foreseeable time. agree with you that the Service should plan an orderly retirement Ecology, as being study of marine organisms and their environ- from sardine research, beginning immediately with the preparation ment, had been immeasurably large, and even study of the rela- and publication of findings.” (Memorandum, Walford to Kask, tiom between organisms and their environment that determine

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their distribution and abundance was proving too large. How 21. Revelle to Isaacs, Nov. 29, 1947, SI0 Director’s Files. could the problem be effectively narrowed? Obvious narrowing 22. It should be noted that whereas Revelle stressed the anomalous was to take one or a few organisms and one or a few local envi- situation that prevailed, as a cause of sardine depletion, other sci- ronments. Study of an organism throughout its range in distri- entists stressed that it was the “normal conditions” or “average bution seemed advisable in order to see the picture through conditions” which had to be identified - that is, “normal” relation- contrasts. . . . ships in the ecological system within which the sardines existed 1.3. There was also a remarkable sense of shared excitement. For as (Walford 1948). Roger Revelle once recalled, in that era of ocean science, since etwy The rhetoric makes it seem, at first blush, that the conceptions in thing needed to be studied, virtually every expedition was certain to question were at odds. But I think that however they phrased the turn up important new data, every plankton-net haul brought up problem rhetorically, the principal designers of the New Oceanog- surprises (Sharp 1988; Revelle 1986). raphy’s approach to ecological systems-both for the tuna, in the 14. In May 1945, for example, Frances Clark of the California fisheries POFI project, and for the sardine, in the MRC project- recognized laboratory set forth her reflections on what could be learned from that “normal” relationships had to be defined in order to understand the catch analyses that her agency had been doing since the 1920s - what deviations from those norms, or anomalies, affected repro- and what problems remained, apparently beyond what catch data duction, survival, abundance, and availability of the species. The and tagging could illuminate. “Boat catch studies,” she observed, problem was dealt with in a revealing letter by Sette, discussed in “will tell us if the sardine population is holding its own, gaining or text at note 24 below. loosing [sic] as the result of fishing. It will not explain changes 23. Revelle to Isaacs, Nov. 29,1947, SI0 Director’s Files, SI0Archives. which occur.” Proposing closer studies of age groupings in a sam- 24. Sette to J. G. Burnette, Nov. 15, 1946, Fish and Wildlife Service pling program to complement the boat catch data, Clark observed Records, File 829.1, Record Group 22, U. S. National Archives. that “the weakest link in our whole investigation is our lack of Although not expressing the vision of ecosystem research so knowledge of recruitment.” She proposed that the sardine investi- explicitly as did Revelle or Sette, R. E. Foerster, director of the gations should thus be expanded significantly, to include both Pacific Biological Station of the Fisheries Research Board of Can- surveys of young fish and larval fish surveys, and “general ada, similarly anticipated a research design for marine fisheries oceanographic investigations.” (Letter from Frances Clark to Rich- studies that would seek to isolate the relevant variables in physical ard Van Cleve, May 1945, Van Cleve Papers, UW Archives.) environment: “It has seemed to me,” he wrote in 1948, Another illuminating exchange between these two sardine ex- that in tackling the biological phase of oceanography - and it is perts, four years later, dealt with the importance ofjuvenile survival an important phase in developing the general picture of the rela- and what was needed technically to do the necessary kind of re- tion of variation in oceanographic conditions to variations in search (letter of Dec. 29,1949, from Van Cleve to Clark, in the Van abundance and/or availability of fish populations - we should, Cleve Papers, UW Archives). for the first few years at least, explore the importance of many The relationship of upwelling to nutriment levels, and the latter factors, such as variations in nitrates, phosphates, carbonates, in relation to spawning, had been opened up for the sardine in the oxygen, phytoplankton, zooplankton[;] determine the relation- California Current by research conducted by Sverdrup on upwell- ships, if any, with a view to subsequently eliminating as many ing and then specifically by Ahlstrom on salinity patterns and as possible and retaining for general survey only those that seem spawning, in 1946-47. This research and its implication are dis- to have a real bearing or influence on abundance or availability of cussed in a memorandum by Harald Sverdrup (1948). fish and can be used for prediction purposes, if such is ever In 1949, Frances Clark was excitedly engaged in preparing a feasible. There are obviously limits to how much field work and paper on the management of pelagic fisheries, hoping to “develop collection of samples, etc., can be done by a vessel and its tech- the need for sound biological, statistical, and oceanographical in- nical and scientific personnel. . . . formation, and thorough cooperation between fisheries investiga- tors in the entire Pacific area.” (Manuscript letter, 1949, in the Van Foerster to J. L. McHugh, Aug. 16, 1948, in Subject Files: Marine Cleve Papers.) For Sverdrup and associates on upwelling, see also Life Research Program, SI0 Archives. (Foerster at this time was Sverdrup et al. 1942. preparing plans for the Nanaimo-based oceanographic project 15. Biographical data on these figures is scattered throughout Shor SARDINE, and was in correspondence with the California group 1978, Scheiber 1986, McEvoy 1986, and Sharp 1988. concerning possible coordination. See Dale Leipper [SIO] to John 16. “Memorandum on the Need for Oceanographic Studies for Pacific P. Tully, Aug. 10, 1948, Subject Files: Marine Life Research Pro- Coast Fisheries,” Manuscript, marked 9 Oct. 1946 (signed by Jo- gram, SI0 Archives.) seph Craig, Frances N. Clark, Donald McKernan, and Oscar E. 25. Indeed Chapman in particular, playing the parlous role of middle- Sette), copy in Director’s Files, SI0 Archives. man between the industry and the scientists, repeatedly warned 17. R. 0. Clover (Navy Hydrographic Office) to Albert M. Day (Fish that “research on the high seas is expensive and time consuming,” and Wildlife Service), Feb. 11, 1947, copy in SI0 Director’s Files, and that industry needs the “damned biologists,” like it or not. SI0 Archives. (Letter to Phister, May 2, 1947, copy in William F. Thompson Pa- 18. Iselin, remarks to the conference “The Position of SI0 in the Uni- pers, UW Archives, Seattle.) versity, the State, and the Nation” (La Jolla, March 1951), transcript 26. Letccr of Chapman to Miller Freeman, Aug. 11, 1947, Miller Free- (copy in SI0 Archives). man Papers, UW Library. On the same lines, Chapman constantly reiterated the theme that 27. The Technical Committee was composed of Robert Miller of the the sardine project was only one strand in a “web of research” that California Academy, Sette of the federal service, Sverdrup, and embraced the entire Pacific Ocean (see Chapman 1947). Richard S. Croker of the California Division of Fish and Game 19. Revelle, remarks to the conference “The Position of SI0 in the (head ofthe Marine Fisheries Research Laboratory and its studies at University, the State, and the Nation” (La Jolla, March 1951), tran- sea). (Minutes ofthe MRC, April 28, 1948, in SI0Archives.) script (copy in SI0 Archives). 28. Sverdrup and Walford had collaborated in studies of upwelling in 20. Letter of Revelle (Office of Naval Research, Washington) to Col. relation to sardine spawning, and Walford had continued his larvae I. M. Isaacs (California Sardine Products Institute), Nov. 29, 1947, and egg studies in 1946-47 in waters off Point Conception and Baja copy in SI0 Director’s Files, SI0 Archives. Revelle incorporated California. The work is described in a letter by Sverdrup (1948). verbatim some of these same passages in the early proceedings of 29. Minutes of the MRC, April 28 and May 19, 1948, SI0 Archives; CalCOFI, in a presentation of the projected SI0 role in the coop- Sette to Revelle, May 11, 1948, Marine Life Research Files, SI0 erative project Memorandum: Marine Life Research Program, May Archives. 3, 1948, manuscript in Subject Files: Marine Life Research Pro- 30. Miller report, in April 1948 MRC Minutes, SI0 Archives. The gram, SI0Archives. original manuscript has notations (in John Isaacs’ hand?) indicating SCHEIBER: CALCOFI‘S EARLY YEARS CalCOFl Rep., Vol. 31,1990

the agency to which each function was primarily assigned (shown SI0 Archives.) Here, then, was the formal origin of the annual in bracketed comments in extract quoted in text, above). CalCOFI Reports that in 1990 recognize the project’s fortieth 31. John Marr (acting chief, Southern Pacific Investigations, US Fish anniversary. and Wildlife Service) to Carl Eckart (director, SIO), Sept. 1, 1948, 49. Hubbs, discussing Ahlstrom’s views, in letter of Hubbs to Chan- SI0 Director’s Files, SI0 Archives; Report of a conference between dler Harris (UCLA Public Information Office), 3 March 1952, in Walford, Silliman. Marr, Eckart, and Revelle, Washington, D. C. Marine fife Research: Publicity file, Hubbs Papers, SI0 Archives. 9-17-48 (manuscript), in SI0 Subject Files: Marine Research Com- 50. University of California, Public Information- Radio, “The Miss- mittee, SI0 Archives; MRC Minutes, Sept. 27, 1948 (includes ing Sardine,” Broadcast #3061, U.E. 1260, Sunday, April 6, 1952, budget items), Subject Files: MRC, SI0 Archives. Columbia Broadcasting System, Los Angeles (manuscript radio 32. Memorandum of Marine Life Research Conference, Dec. 30, 1948 text, copy in Hubbs Papers, SI0 Archives). (manuscript dated Jan. 10, 1949), SI0 Subject Files: Marine Life 51. That is to say, I still adhere to the view taken in McEvoy and Research, SI0 Archives. Scheiber 1984 (page 406), but which my coauthor in that study has 33. Eckart to Walford, June 28, 1948, SI0 Director’s Files: Marine Life largely abandoned (see McEvoy 1986), that “the very complexity of Research, SI0 Archives. ecology research - rendered progressively more complex by the 34. San Diego Union, Nov. 28, 1948, clipping in Carl Hubbs Papers, emerging interdisciplinary approach that MRC funds fostered- SI0 Archives (quoting testimony of Richard Croker before the made delay and indecision on policy a more likely result, at least for state assembly’s interim committee on fish and game, citing the several years.” It was probably the scientists’ concern “not to hurry drop in the catch from 403,700 tons in 1945-46 to 124,200 tons in or be popular, but to be right. . . . That the resultant stalemate 1948; Croker recommended a 100,000-ton limit). played into the hands of an industry that wished to avoid regulation 35. Revelle to John Marr, Dec. 14, 1948, copy in SI0 Subject Files: was in that respect incidental - though it had tragic consequences Marine Life Research, SI0 Archives. (Marr had suggested that the for the fishery” (McEvoy and Scheiber 1984). cruise of the federal ship Black Doriglax be postponed until the See also the views of Radovich (1981), stressing “agency-based April-July period, to permit full reoutfitting. Marr to Eckart, Nov. perspectives” that he feels led the state scientists (committed to 22, 1948, ibid.) regulation) in a direction divergent from that which the entire cor- 36. Marine Life Research Conference, memorandum of Dec. 30, 1948, porate history of their agency suggested was the best, or at least the meeting with participating agencies (dated Jan. 10, 1949), copy in prudent, course for the federal scientists. SI0 Subject Files: Marine Life Research, SI0 Archives. Years later, some of the leading scientists who had been associ- 37. Standard analyses of federal-state relations, in the literature of fed- ated with CalCOFI since its beginnings explicitly voiced this view eralism and governance, of course treat at length the various types ofthe need for neutrality. Thus Revelle and John Isaacs, responding of federal grants-in-aid to states but entirely neglect even the pos- to pressures for the CalCOFI scientists and the MRC to take a sibility that the flow might ever run in the opposite direction. See, position on a key matter of policy regarding anchovy reduction e.g., Wright 1982; cf Scheiber 1980. plant permits, warned “that the MRC and CalCOFI should remain 38. Plans ofthe Bureau of Marine Fisheries, California Division of Fish non-political and should not enter into the existing [policy] contro- and Game, for Expanded Sardine Research and Budget Requests of versies.” The chairman of MRC since its founding, Robert Miller the Marine Research Committee (manuscript marked “July 20, of the California Academy, then “read from section 729 of the Fish 1949”), copy in SI0 Subject Files: Marine Life Research, SI0 and Game Code which . . . essentially [read] that MRC cannot Archives. make recommendations, it can only point out facts and make esti- 39. Ibid. (On the continuing political travails of the state agency’s sci- mates of the situation.” (Minutes of MRC meeting of Aug. 13, entists and their efforts to bring the sardine fishing under control, 1963, copy in Subject Files: Marine Research Committee, SI0 Ar- see McEvoy 1986.) chives.) Later on, Miller wrote of the “superb job of getting previ- 40. Progress report of the South Pacific Fishery Investigations, U.S. ously warring agencies to work peaceably and even enthusiastically Fish and Wildlife Service, in the Expanded Pilchard Research Pro- together” as an important achievement ofCalC0FI and basis for its gram, 1 May-31July 1949 (manuscript report), copy in SI0Subject research accomplishments. (Robert Miller to Wilbert Chapman, Files: Marine Life Research Program, SI0 Archives. Feb. 3,1964, Robert Miller Papers, California Academy of Sciences 41. Ibid. Archives.) 42. Roger Revelle to Robert C. Miller, Sept. 13, 1949, enclosing copy Also relevant in coming to a judgment of scientists’ behavior in of the May 1-July 31, 1949 SI0 report, copy in SI0 Subject Files: this era is the commitment of some, such as Ahlstrom, that the Marine Life Research Program, SI0 Archives. “extremely important function” of MRC as “one of the best coor- 43. Ibid. dinating mechanisms he knew of in fisheries research . . . has kept 44. Letter of Clark to Hubbs, June 1, 1950, Subject Files: Marine Life people working amicably in the same ocean on the same problem;” Research, SI0 Archives. and that any split caused by dealing with explosive political issues 45. Ibid. See also McEvoy 1986, pp. 200-201, for more substantive that could be resolved in other arenas would work against this controversy between the state agency and the federal and SI0 sci- coordination, which “he submitted. . . (was) the greatest value of entists, concerning the proper way in which the issue of sardine MRC and, . . should be preserved.” (Minutes of MRC meeting of depletion vis-i-vis commercial fishing ought to be presented to the Jan. 19, 1965, SI0 Archives.) public. Resolution of the difference in interpreting the scientists’ and 46. Ahlstrom 1950, italics added. MRC roles must turn, at least in part, on whether one judges that 47. Letters from Marr to Revelle, Sept. 9, 1949; Clark to Marr, Sept. the evidence of harm to stocks from overfishing was so compelling 13, 1949; and Revelle to Marr, Sept. 12, 1949, copies in SI0 Subject by even 1947, let alone 1952, that scientists who failed to register Files: Marine Life Research Program, SI0 Archives. opinions on the side of suspension or tighter regulation were In 48. Annual reporting and publication of the scientific projects (aug- effect irresponsible. See also text, infra, at note 57. menting the quarterly agency reports) were ordered beginning in 52. Especially so, of course, by dint of the intensive (monthly) data 1950, after discussion at theJune 7-8, 1950, meetings of the Tech- collection at all stations of the enormous grid that was established nical Advisory Committee and the MRC. Such a report, the coni- in 1948-49. (See CalCOFI 1989; Revelle 1986.) A few years later, an niitters declared, would serve as “a summary of progress and MRC member wrote that “Our [California] offshore seas and their results to date. . . . [and] should be widely distributed . . . as a inhabitants are better known and understood than any in the world basis for consideration by the fishing industry and the legislature of with the possible exception of the Norwegian Sea” (Bruce 1963). In the desirability of continuing the program of the Marine Research 1959John Isaacs asserted, “It is safe to say that there has never been Committee.” (Revelle Memorandum, July 13, 1950, to Thomas another study that resulted in so thorough an understanding of a Manar, copy in Marine Life Research: Publicity file, Hubbs Papers, pelagic species of fish as that [which] CalCOFI and earlier studies SCHEIBER: CALCOFI‘S EARLY YEARS CalCOFl Rep., Vol. 31,1990

have obtained on the sardine.” (Isaacs, in Appendix to Minutes of in the project. (Chapman, comments reported in Minutes of MRC the Marine Research Committee, July 30, 1959 meeting, SI0 meeting of Jan. 1965, SI0 Archives. See also Scheiber 1986 on Archives.) Chapman’s efforts in this period to liberalize more generally the 53. Four or more spawning areas were early identified, one in the Gulf regulation of California commercial fisheries and to reduce the in- of California, others off southern Baja California, central Baja Cal- fluence of the sports interests.) Murphy’s scholarly contributions ifornia, and an area off the southern California and northern Baja are considered in Ahlstrom and Radovich 1970 and in McEvoy California coast. (Technical Committee report, in MRC Minutes 1986. for Dec. 19, 1957, SI0 Archives.) 65. The political story is recounted in McEvoy 1986, pp. 215-20; on 54. Ibid. Chapman and the industry’s role in the controversy, see Scheiber 55. Report of the Special Technical Committee, MRC Minutes, Dec. 1986. Beyond the purview of the present study is the further re- 19, 1957, SI0 Archives, also quoted in Murphy 1960. It is notewor- search done by Isaacs on sedimentary evidence of the historic sar- thy, also, that indicating the legislature’s (and the fishing industry’s) dine “is an unusual event” in what he termed the normal “hake- recognition and approval of this expanding agenda, new taxes were anchovy complex” of the California Current biomass. (Paper pre- levied on mackerel and anchovy, to augment the revenues (which sented to the MRC meeting of May 11, 1965, Minutes, SI0 Ar- were steadily declining because of the continued fall in sardine chives.) On the various studies by Lenarz, Smith, and McCall on landings) from the original sardine tax authorized in 1947. State and the interpretation of the anchovy-sardine data and the implications federal general appropriations for SIO, USFWS, and California for management that this important work suggested, see the dis- state agency research continued to support the larger program that cussion in McEvoy 1986, pp. 232-235. the MRC funds augmented, so that in 1959 it was estimated that the total spent for programs directly linked to CalCOFI research represented a level of $130,000 of MRC funds from the landings REFERENCES taxes, $600,000 of SI0 funds, $250,000 of USFWS funds, and Ahlstrom, E. H. 1950. A report on the problems and objectives of the $200,000 of the California Department of Fish and Game funds. cooperative sardine research program [marked “May 1950,” written (MRC Minutes ofDec. 1,1959 meeting, SI0 Archives.) with assistance from persons working in the program], copy in SI0 56. Report of the Special Technical Committee, Minutes of MRC Subject Files: Marine Life Research: General, SI0 Archives. meeting, Dec. 19, 1957, SI0 Archives. -. 1963. Larval fish studies (manuscript report, prepared for Ma- 57. Reference here is to the work of Ahlstrom, Isaacs, Murphy, and rine Research Committee), in Minutes of MRC meeting of ll April Paul Smith in the post-1960 period as well as to that of Ahlstrom, 1963, copy in Subject Files: MRC, in SI0 Archives. Walford, Marr, and Clark in the years from 1937 to 1960. On their -. 1964. Egg and larval surveys (manuscript report, prepared for respective contributions, see, inter alia, McEvoy 1986, Ahlstrom Marine Research Committee), in Minutes of MRC meeting of 6 and Radovich 1970. Throughout the entire period of CalCOFI re- March 1964, Appendix, copy in Subject Files: MRC, in SI0 search to the mid-1960s. the California Fish and Game scientists Archives. unsuccessfully sought urgently to obtain full regulatory powers Ahlstrom, E. H., and J. Radovich. 1970. Management of the Pacific over the sardine fleet, but even the definitive collapse that occurred Sardine. In A century of fisheries, N. Benson, ed. American Fisheries in 1952-53 (when the catch went from 145,000 tons to 15,000) failed Society, Special Publication No. 7, Washington: American Fisheries to win them the authority they sought (Ahlstrom and Radovich Society, pp. 183-193. 1970). Barrett, I. 1980. Development of a management regime for the Eastern 58. The great weather shift that occurred in 1957 and 1958 did finally Pacific tuna fishery. Ph.D. dissertation, College of Fisheries, Univer- give the SI0 and other CalCOFI scientists new insight into varia- sity of Washington, Seattle. tions from normal conditions and their impact on the fisheries. A Baxter, J. L. 1982. The role of the Marine Research Committee and major symposium was held in 1958- “1957 and 1958, the Years of CalCOFI. Calif. Coop. Oceanic Fish. Invest. Report 23:35-38. Change”-and reported on by John Isaacs in Minutes of MRC Bruce, E. F. 1963. The impact of new technology on ocean fish conser- meeting, June 10,1958, SI0 Archives. vation and management as seen by the Marine Research Committee 59. Letter of Sette to J. G. Burnette, Chairman, MRC, Dec. 4, 1957, in (transcript of talk before the Wildlife Society, no date), copy in Min- Minutes of MRC meeting, Dec. 19, 1957, SI0Archives. utes of MRC meeting of March 1963, Subject Files: MRC, SI0 60. McKernan to Burnette, Dec. 12, 1957, ibid; 0. E. Sette to Burnette, Archives. Dec. 4, 1957, ibid. CalCOFI. [1989] California Cooperative Oceanic Fisheries Investiga- 61 Report of the Special Technical Committee, in Minutes of MRC tions. (CalCOFI, no date, but 1989), not paginated. Meeting, Dec. 19,1957, SI0 Archives. Chapman, W. M. 1947. Statement in testimony, in 80th Cong., 1st Sess., 62 Letter of Chapman to Phister, Sept. 19, 1947, Chapman Papers, House of Rep. Comm. on Merchant Marine and Fisheries, Develop- UW. Chapman had wanted John Kask, then ofthe California Acad- ment ofthe high seas fishing industry: Hearings, May 12,1947. Wash- emy of Sciences, to be named coordinator, partially because of ington: GPO, 1949, pp. 3-11. Kask’s personal qualities but partially too because the Academy Deacon, M. 1990. The establishment of marine stations and early com- was seen as a neutral player in the politics of fisheries science in mercial fisheries research in Scotland. Unpublished paper for Califor- California. As it worked out, the Technical Advisory Committee nia Sea Grant College Program/UC Intercampus Economic History that was appointed in 1948 served the coordinating function, and program conference, Berkeley, May 1990. Copy on file, Center for Director Robert C. Miller of the Academy was its chairman for the Study of Law and Society, UC Berkeley. more than 17 years. (Robert C. Miller to Chapman, Feb. 3, 1964, Dymond, J. 1948. European studies of the populations of marine fish- Robert C. Miller Papers, California Academy of Sciences eries, Bulletin of the Bingham Oceanographical Collections 11:55. Archives.) Fleming, D. 1971. Roots of the new conservation movement. Perspect. 63 The quotation is from Miller to Chapman, Feb. 3, 1964, Miller Am. Hist. 6:7-91. Papers, California Academy of Sciences Archives. Herre, A. 1940. Distribution of fish in the tropical Pacific. In Sixth 64 See Baxter 1982. Later, in the mid-l960s, when MRC had been Pacific Science Congress, Proceedings, Pacific Science Academy expanded to include sportfishing and labor representatives, and 3:587. when the frustration of the commercial fishing and cannery inter- Herrington, W. C. 1988. Interview by the author, Staffordville, Con- ests with a standoff-in MRC and in state policy bodies-on pro- necticut, June 1988. (Notes in author’s files.) posals to open and expand the anchovy fishery caused new and -, 1989. In the realm of diplomacy and fish: some reflections on deeper rifts within MRC, the coordinator did come under some International Convention on High Seas Fisheries in the North Pacific heavy criticism for what one industry representative (Chapman) Ocean and the Law of the Sea negotiations. Ecology Law Quar. regarded as his failure to exercise sufficient control over the agencies 16:1087. SCHEIBER: CALCOFI‘S EARLY YEARS CalCOFl Rep., Vol. 31,1990

Huntsman, D. 1949. Fisheries oceanography (manuscript received in Scheiber, H. N. 1980. Federalism and legal process: historical and con- May 1949 by the SI0 Director’s Office and circulated at SIO), in SI0 temporary analysis of the federal system. Law and Society Rev. Subject Files: Marine Life Research, SI0 Archives. 14:663-722. Idyll, C. P. 1969. The science of the sea. 1iz Exploring the ocean world: -, 1986. Pacific Ocean resources, science, and Law of the Sea: a history ofoceanography. New York: Crowell, pp. 2-21. Wilbert M. Chapman and the Pacific Fisheries, 1945-70, Ecology Isaacs, J. 1959. Proposed program in fisheries research (manuscript re- Law Quar. 13:381-534. port to Marine Research Committee), in Minutes of MRC meeting -. 1988. Wilbert Chapman and the revolution in U.S. Pacific of 30 July, 1959, copy in SI0 Subject Files: MRC, SI0Archives. Ocean science and policy, 1945-1951. bi Nature in its greatest extent: Knauss, J. 1990. Remarks at Symposium of CalCOFI Conference, Oc- western science in the Pacific, E? Rehbock and R. MacLeod, eds. tober 27, 1989. Calif. Coop. Oceanic Fish. Invest. Rep. 31: (this Honolulu: University of Hawaii Press. volume). -. 1989. Origins of the Abstention Doctrine in ocean law: Japa- Larkin, P. A. 1970. Management of Pacific salmon of North America. nese-U.S. relations and the Pacific fisheries, 1937-1958. Ecology Law In A century offisheries, N. Benson, ed. American Fisheries Society, Quar. 1623-99. Special Publication No. 7, Washington: American Fisheries Society, . In press, a. U.S. ocean fishery research in the Pacific, 1945- pp. 223-235. 1970, Deutsche Hydrograph. Zeit. Los Angeles Times. 1949. Fleet of six ships launches biggest fish hunt in . In press, b. Science, commercial interests, and the policy pro- history. Los Angeles Times, June 22, 1949, clipping in Publicity: cess in U. s. oceans research: the POFI tuna program, 1948-64. Paper MLR file, Carl Hubbs Papers, SI0Archives. to the International History of Science Congress, Hamburg and McEvoy, A. 1986. The fisherman’s problem: ecology and law in the Munich, to be distributed as an Ocean Law and Policy Working California fisheries, 1850-1980. New York: Cambridge University Paper, Center for the Study of Law and Society, School of Law, Press. University of California, Berkeley. McEvoy, A., and H. N. Scheiber. 1984. Scientists, entrepreneurs, and Sette, 0. E. 1943. Studies on the Pacific pilchard or sardine (Savdirzops the policy process: a study of the post-1945 California sardine deple- caevulea):I. Structure of a research program to determine how fishing tion. J. Econ. Hist. 44:393-413. affects the resource. U.S. Fishand Wildlife Service, Spec. Sci. Report, McHugh, J. L. 1970. Trends in fishery research. Iiz A century of fisher- No. 19. ies, N. Benson, ed. American Fisheries Society, Special Publication . 1952. Letter to POFI industry advisory committee, June 20, No. 7, Washington: American Fisheries Society, pp. 25-56. 1952, Pacific Oceanic Fishery Investigations Records, U. S. Fish and Miles, E. A,, S. Gibbs, D. Huhart, C. Dawson, and D. Teeter. 1982. Wildlife Service Papers, National Archives. The management of marine regions: the North Pacific: an analysis of -. 1955. Considerations of mid-ocean fish production as related issues relating to fisheries, marine transportation, marine scientific to oceanic circulatory systems. J. Mar. Res., 14:398. research, and multiple use conditions and conflicts. Berkeley: Univer- Sette, 0. E., and E. H. Ahlstrom. 1948. Estimations of the abundance sity of California Press. of the eggs of the Pacific pilchard (Savdinops cawulea) off southern Murphy, G. 1960. Operational analysis of CalCOFI research (Sept. 7, California during 1940 and 1941. J. Mar. Res. 7:511-541. 1960). Report to Marine Research Committee, in Minutes of MRC Sharp, S. 1988. Observations on the Office of Naval Research and inter- meeting, 4 Oct. 1960, Exhibit #1, copy in SI0 Subject Files: MRC, national science, 1945-1960: oral history interview of Roger Revelle. SI0 Archives. La Jolla: Scripps Institution of Oceanography. -. 1963. Summation of Calcofi. (Manuscript report to Marine Shor, E. N. 1978. Scripps Institution of Oceanography: probing the Research Committee), in Minutes of MRC meeting, 11 April 1963, oceans, 1936 to 1976. San Diego: Tofua Press. copy in Subject Files: MRC, SI0 Archives. Sverdrup, H. U. 1948. The sardine problem. Manuscript memorandum Murphy, G., and J. Isaacs. 1964. Species replacement in marine ecosys- prepared for a journal editor as a background document and enclosed tems with reference to the California Current. (Manuscript report), with a letter by Sverdrup to Charles M. Hatcher (editor, Tuna Fish- in Minutes of meeting of MRC, March 6, 1964, Appendix, copy in erman), Feb. 23,1948, SI0 Subject Files: MRC, SI0 Archives. Subject Files: MRC, SI0 Archives. ,1952. Some aspects ofthe primary productivity ofthe sea. FA0 Pacific Fisherman. 1947. August 1947 symposium issue (special sym- Fish. Bull. 5(6):215-223. posium under the title, “Who Will Harvest the Pacific,” introduced Sverdrup, H. U., M. W. Johnson, and R. H. Fleming. 1942. The oceans: by the editors as an “outline of the resources latent in the Pacific their physics, chemistry, and general biology. New York: Prentice- Fisheries Frontier”). Hall. Pan-American Fisherman. August 1950. (Summary of SI0 role in sar- Thompson, W. E 1919. The scientific investigation of marine fisheries, dine research and related oceanographic studies) Clipping in Subject as related to the work of the Fish and Game Commission in southern Files: Marine Life Research, SI0 Archives, p. 4. California. California Fish and Game Commission Fish. Bull. No. 2. Radovich, J. 1981. The collapse of the California sardine fishery. In Walford, L. A. 1948. The case for studying normal patterns in fishery Resource management and environmental uncertainty, M. H. Glantz biology, J. Mar. Res. 7:506-510. andJ. D. Thompson, eds. New York: Wiley, 110-111. Wooster, W. 1949. Memorandum, Warren Wooster to Roger Revelle, 3 Revelle, R. R. 1986. Interview by the author, La Jolla, May 1986. Notes May 1949, Subj: Personnel assigned to MLR chemistry division, Ma- in author’s files. rine Life Research files, SI0 Archives. Russell, E. S. 1942. The overfishing problem. The De Lamar Lectures Wright, D. S. 1982. Understanding intergovernmental relations, 2nd at the Johns Hopkins University. Cambridge: Cambridge Univ. edition. Montery, Calif.: Brooks Cole. Press.

83 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

SETTLEMENT OF JUVENILE CALIFORNIA HALIBUT, PARALICHTHYS CALIFORNICUS, ALONG THE COASTS OF LOS ANGELES, ORANGE, AND SAN DIEGO COUNTIES IN 1989

M. JAMES ALLEN KEVIN T. HERBINSON MBC Applied Environmental Sciences Southern California Edison Company 947 Newhall Street F? 0. Box 800 Costa Mesa, California 92627 Rosemead, California 91770

ABSTRACT disefiado con el prop6sito de: (1) determinar el re- Juvenile California halibut (Pavulichthys calijovni- clutamiento relativo de 10s juveniles, y (2) determi- cus) typically occur in bay nursery grounds along the nar la supervivencia de 10s juveniles pequefios en las coast of California and the west coast of Baja Cali- zonas poco profundas. Se estudiaron algunas 5reas fornia. Recent surveys using small-meshed beam costeras de 10s condados de Los Angeles, Orange y trawls indicate that some settlement to the bottom San Diego, de abril a septiembre de 1989. Se toma- also occurs in shallow coastal areas of southern Cal- ron muestras en cuatro localidades costeras (Her- ifornia. This survey of halibut settlement in bay and mosa Beach, Long Beach, San Onofre y Carlsbad) coastal areas was designed to determine (1) the rela- y en dos lagunas costeras (Anaheim Bay y Agua tive settlement rates of juvenile halibut in bay and Hedionda Lagoon). Se muestrearon tres estaciones coastal environments and (2) the survival of small en cada localidad utilizando una red de arrastre de juveniles in the shallow coastal zone. Areas off Los bao de 1.0 m en las lagunas y una red de 1.6 m en las Angeles, Orange, and San Diego counties were sur- Preas costeras. Se colectaron 288 muestras en total, a veyed from April to September 1989. Three stations profundidades de 0-3 m en las lagunas y de 6-13 m were sampled at each of four coastal sites (Hermosa a lo largo de la costa. El estudio de estas muestras Beach, Long Beach, San Onofre, and Carlsbad) and indic6 que en 1989 10s juveniles del lenguado de two bay sites (Anaheim Bay and Agua Hedionda California se reclutaron en las aguas costeras poco Lagoon). A 1.0-m beam trawl was used in bays and profundas de 10s condados investigados. El mayor a 1.6-m beam trawl along the coast. A total of 288 reclutamiento se present6 en Anaheim Bay, Her- samples was collected at depths of 0-3 m in bays and mosa Beach y Long Beach. Los juveniles permane- 6-13 m along the coast. These samples indicated that cieron en las bahias costeras durante el period0 de juvenile California halibut did settle into shallow estudio. Si bien tambitn se encontraron algunos in- coastal waters of the areas surveyed during 1989. dividuos en otros sitios costeros durante el estudio, Settlement was greatest at Anaheim Bay, Hermosa el reclutamiento exitoso occurri6 solamente de julio Beach, and Long Beach. The settling halibut re- a septiembre y unicamente en 10s sitios semiprote- mained in bays throughout the study period. Al- gidos (Hermosa Beach y Long Beach). though transforming fish were also found at coastal sites throughout the study period, successful settle- INTRO DUCTlO N ment occurred only from July to September and California halibut (Pmalichthys cal$ofornicus) is an then only in the semiprotected sites (Hermosa Beach important species to the ecology and fisheries of and Long Beach). coastal southern California. As ajuvenile it is a char- acteristic component of the bay (i.e., coastal lagoon) RESUME N fish community. As an adult it is an important am- Los juveniles del lenguado de California (Pudich- bushing predator of nearshore fishes (M.J. Allen thys calfofovnicur) se encuentran en general en las bahias 1982) and an important species in the marine sport criaderas a lo largo de la costa de California y de la and commercial fisheries of California (NMFS 1985; costa occidental de la peninsula de Baja California. CDFG 1989). Estudios recientes con red de arrastre de bao de Crucial portions of habitat are being threatened malla fina indican que el establecimiento en el fondo by human encroachment. California halibut are se da en ciertas Preas costeras de poca profundidad generally thought to require bays for nursery del sur de California. Este estudio del estableci- grounds, and thus these areas may be crucial to their miento del lenguado en bahias y ireas costeras fue survival (Haaker 1975; Kramer and Hunter 1987; L. G. Allen 1988a; Kramer, in press). Most small ju- ~ ~____ [Manuscript received February 1,1990.1 veniles are found in bays; it is only at a larger size

84 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

that halibut move to coastal waters, where they re- (Anaheim Bay and Agua Hedionda Lagoon), two cruit to the fisheries (Haaker 1975). semiprotected coastal sites (Hermosa Beach and Until recently, little was known about the settle- Long Beach), and two exposed coastal sites (San ment of California halibut from the plankton to the Onofre and Carlsbad). The Agua Hedionda Lagoon, bottom. The abundance ofjuveniles in bays has been Carlsbad, and San Onofre sites are located in San known for some time (Haaker 1975), as has the pau- Diego County and have been sampled in previous city of small juveniles (<150 mm) in open coastal surveys (Kramer and Hunter 1987, 1988). Anaheim areas (M.J. Allen 1982; Plummer et al. 1983). How- Bay is in Orange County; Long Beach and Hermosa ever, the distribution and biology of newly settled Beach are in Los Angeles County. juveniles has only recently been studied (Kramer The bay sites are fully protected from offshore and Hunter 1987; L.G. Allen 1988a, b; Kramer and swells; the semiprotected coastal areas are partially Hunter 1988; L.G. Allen et al., in press; Kramer, in protected; and the exposed coastal sites are more press). fully exposed to offshore swells. Depending on Surveys in the vicinity of Alamitos Bay in 1983- wind and swell direction, the Hermosa Beach site 85 indicated that recently settled California halibut may be variably exposed or protected during the were most abundant in the protected habitat of year. It is protected from swells from the south or Alamitos Bay, less abundant in the semiprotected southwest during the summer by the Palos Verdes habitat of Long Beach Harbor, and least abundant Peninsula and offshore islands but is exposed to on the open coast at Sunset Beach (L.G. Allen western swells, which occur primarily during the 1988a). Recent surveys in bays and open coast areas winter (Maloney and Chan 1974). The Long Beach of San Diego County revealed little coastal settle- site is protected from swells from the northwest, ment in 1987, but substantial coastal settlement in west, and southwest by the Palos Verdes Peninsula, 1988 (Kramer and Hunter 1987, 1988; Kramer, in breakwaters, and offshore islands. The San Onofre press). Another survey in 1988 along the open, near- site is fully exposed to swells from the south and the shore coast of southern California from Point Con- southwest, and the Carlsbad site to swells from the ception, Santa Barbara County, to San Mateo Point, west and south. Orange County, found that the greatest numbers of At all sites, stations were located randomly within settling individuals occurred in southern Santa blocks stratified by depth. The water depth of the Monica Bay and in Long Beach Harbor (L.G. Allen blocks ranged from 0.0 to 0.8 m, 1.0 to 1.5 m, and et al., in press). 3.0 to 3.5 m in bays, and from 6 to 8 m, 8 to 11 m, Thus, although settlement of California halibut and 11 to 15 m along the coast. For analyses, these to the open coast of southern California has been blocks are represented as stations with depths of0.5, described, little is known of halibut's interannual 1.0, and 3.0 m in the bays and 6.0, 10.0, and 13.0 m settlement success there. Because of the importance of California halibut to the fisheries, and the contin- uing encroachment of human activity in the bay nursery grounds, it is important to continue studies of settlement patterns. The objective of this study was to determine the relative settlement and survival of juvenile Califor- nia halibut in selected bay and coastal environments of Los Angeles, Orange, and San Diego counties during the spring and summer of 1989.

METHODS

Study Area The study area extended from Hermosa Beach to Carlsbad, California. Within this area six sites were surveyed: Hermosa Beach, Long Beach, Anaheim

Bay, San Onofre, Carlsbad, and Agua Hedionda La- Figure 1. Locations of beam trawl surveys for juvenile California halibut (Par- goon (figures 1 and 2). These included two bay sites alichthys californicus),April-September 1989.

85 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990 a)

t*HB2 ", \ '* '* kermosa Beach

", 'b \ HB1 \

Long Beach\- h 1 Alamitos Bay ---Mv ruI

AH2 Carlsbad -1 km

\ 2 0 AB3 1 \,,, Agua Hedionda Lagoon -0.5 km 5,

Figure 2. Stations sampled by beam trawl along the southern California coast, April-September 1989: a, Hermosa Beach; b, Long Beach; c, Anaheim Bay; d, San Onofre; e, Carlsbad;and f, Agua Hedionda Lagoon. along the coast, approximately the same depths as were collected with a 1.6-m beam trawl and bay sampled by Kramer and Hunter (1987, 1988). Sta- samples with a 1.0-m beam trawl. All nets had 2.5- tion coordinates are given in MBC 1990. mm stretch-mesh netting. The beam trawls were equipped with a wheel and revolution counter (me- Sampling Methods ter wheel) which recorded the distance that the trawl Fish were collected with the same nets used by traveled on the bottom, although the meter wheel Kramer and Hunter (1987, 1988). Coastal samples occasionally clogged with plant debris. At coastal

86 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

stations loran C coordinates (longitude and latitude) iba analyzer. Station values of these oceanographic were recorded at the beginning and end of each haul. parameters are given in MBC 1990. In the bays, 200-m trawl paths were measured and marked with buoys to provide a separate indication Data Analysis of towing distance, in case the meter wheel fouled. The bottom area actually sampled in each tow was Depth was measured with sonar at coastal sites and calculated using meter-wheel readings or distances with a sounding line at bay sites. measured in the field. When fouling had occurred or Coastal trawling was conducted from the R/V the meter-wheel reading was obviously too low, the Westwind, a 14.6-m research vessel. Bay trawling at distance traveled was estimated. This estimate was 1.0-m and 3.0-m stations was conducted from a 5.2- 200 m in bays and 315 m along the coast. The coastal m Boston whaler. At the 0.5-m stations the beam estimate was based on the average distance attained trawl was pulled by two field workers on foot. by all "good" tows. The area sampled was com- Three 10-min (coast) or 200-m (bay) trawls were puted as the product of the distance towed and the attempted during daylight hours at each site's three width ofthe trawl- 1.0 m for bays and 1.6 m for the stations, for a total of nine replicates per site. This coast. Estimated areas for replicates with extremely should have resulted in 54 samples per survey, but low readings were 200 m' for bays and 504 m' for lost or broken nets and heavy loads of algae often the coast. reduced the available sampling time. Thus fewer Length-frequency histograms of California hali- replicates were completed at some stations. Physical but were generated for each survey and were plotted characteristics of each tow are given in MBC 1990. by area and month. Densities of halibut size groups Each of the six sites was sampled in April, May, at various sites were also determined. The relation- June, July, August, and September of 1989. These ship between settlement and temperature was deter- were the months of major settlement of California mined by linear regression analysis. halibut into the coastal environment off San Diego County during 1988 (Kramer and Hunter 1988), al- RESULTS though settlement into bays can begin as early as November. Although this study emphasized California hali- Sampling Ecfjrort but, all fish captured were retained for identification From April to September, 288 trawl samples were and measurement. Most were returned to the labo- collected, 36 to 52 each month. All stations were ratory for processing; however, large specimens sampled, but fewer than three replicates were ob- were identified to species, measured, weighed, and tained at some stations because of fouling by algae released in the field. Because transforming halibut or sand. Totals of 190 samples were collected along and other juvenile fishes are small, most debris (and the coast and 98 in the bays; of the coastal samples, invertebrates) was returned to the lab for closer ex- 95 were collected at semiprotected and 95 at exposed amination; only large debris was discarded in the stations. From 47 to 50 samples were collected at field. Specimens and debris were fixed in buffered each site, 29 to 35 at each depth in the bays, and 60 10% Formalin-seawater. to 64 at each depth along the coast. In the laboratory the samples were rinsed of For- The total area sampled was 11.5 ha, 1.6-2.1 ha per malin after about a week and transferred to 70% month. A total of9.7 ha was sampled along the coast isopropyl alcohol. Samples were then sorted to sep- and 1.8 ha in bays; along the coast 4.8 ha were sam- arate fish from invertebrates and debris. Fish were pled in the semiprotected habitat and 4.9 in the ex- identified to species, measured, and weighed. The posed habitat. At each coastal site, 2.4-2.5 ha were standard length (SL) ofeach bony fish or total length sampled; at each bay site 0.9 ha were sampled. At of each cartilaginous fish was measured to the near- each depth zone within the bays, 0.6-0.7 ha were est millimeter. For abundant species, subsamples of sampled; at each depth zone along the coast, 3.1-3.5 up to 200 individuals were measured separately. The ha were sampled. total weight of each species in a sample was weighed on a Mettler balance to the nearest 0.01 g. Physical Oceanography Near-bottom water samples were collected at Monthly near-bottom water temperatures at sta- each station with Van Dorn bottles, generally after tions in this study ranged from 13.8" to 24.7"C (table the last haul at the station. Temperature and pH of 1). Monthly temperatures at stations along the coast these samples were measured in the field with a Hor- ranged from 13.8" to 22.0"C and in bays from 18.3"

87 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

TABLE 1 Water Temperature and pH in Coastal and Bay Habitats, 1989

~~ Coastal Month Min. Max. Mean SD Min. ~-~ ~~ ~~ ~~ ~~ - ~- Temperature ("C) April 15 0 18.2 16.4 0.9 18.3 21.7 19.7 1.1 May 13 9 18.3 16.6 1.5 18.3 18.9 18.6 0.3 June 14 4 18.1 16.2 1.3 18.9 22.8 20.9 1.3 July 17 3 22.0 20.1 1.2 21.3 24.7 23.1 1.1 August 13 8 21.1 17.9 2.7 21.0 23.0 22.2 0.9 September 16 1 -30.4 18.0 1.2 20.3 22.8 21.2 0.8

PH April 72 7.7 7.5 0.1 7.3 79 77 02 May 74 7.8 7.7 0.1 7.7 82 80 02 June 66 7.6 7.3 0.3 6.6 69 68 01 July 73 7.9 7.7 0.1 7.9 80 80 01 August 74 10.2 8.1 0.1 7.9 82 81 01 September 79 8.3 8.1 0.1 7.8 81 79 01 ~- ~__~~ ~~ ~ ______~_____

to 24.7"C. Means ranged from 18.6" to 23.1"C in the 30 bays and from 16.2" to 20.1"C along the coast. Monthly mean temperatures at sampling sites were - greater in the bays than along the coast (table 1; 25 figure 3). In the bays the highest mean temperature ?! ca occured in July and the lowest in May. Along the 2 coast the highest mean temperature also occurred in n0) zc July, but the lowest occured in June. The highest I- temperatures in both bay and coastal areas were dur- ing the last three months of the survey period. Hydrogen ion concentration (pH) in this study 1:: ranged from 6.6 to 10.2 (table 1). pH values had a greater range along the coast than in the bays; how- ever, monthly means had a greater range in the bays Figure 3. Mean monthly temperatures at sites surveyed for juvenile California than along the coast. The monthly mean pH values halibut (Paralichthys californicus), 1989. at bay sites were generally equal to or higher than those found at coastal sites, except in June and Sep- bers are a result of variation in sampling effort due tember, when values were higher along the coast. to differences in net sizes used, tow lengths, and Along the coast the lowest mean pH occurred in number of hauls. Standardization of the catch by June and the highest in August and September. In area trawled indicated that the mean halibut density the bays the lowest pH also occurred in June and the was highest (158 fish/ha; SD = 275) in the bays, highest in August. intermediate (94 fish/ha; SD = 184) at semipro- tected coastal sites, and lowest (25 fishlha; SD = 44) Distribution and Settlement at exposed coastal sites. Cutch purunzeters. California halibut occured in The total weight of California halibut taken in the 55% of the samples collected. The fish were almost survey was 37.7 kg, about 15% of the total fish bio- equally common along the coast and in the bays, mass collected (243.3 kg). Total halibut biomass was being found in 56% of the coastal samples and 53% much greater (36.7 kg) at coastal sites than in bays of the bay samples. We collected 762 halibut in the (0.9 kg). Along the coast the biomass was greater survey, about 2% of all the fish captured (48,994). (20.8 kg) in the semiprotected habitat than in the More halibut were taken along the coast (457) than exposed habitat (15.9 kg). Again, these absolute val- in the bays (305). Along the coast, more were taken ues result from differences in sampling effort at bay in the semiprotected habitat (348) than in the ex- and coastal sites. Standardization by area indicated posed habitat (109). However, these absolute num- that the mean biomass density was highest (4.8 kg/

88 ALLEN AND HERBINSON: SElTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

ha; SD = 9.0) in the semiprotected coastal habitat, 30 intermediate (3.3 kg/ha; SD = 6.1) in the exposed 25 coastal habitat, and lowest (0.5 kg/ha; SD = 1.3) in bays. 0 20 Size and population stuuctuve. California halibut ranged in size from 6 to 503 mm SL, with a mean l5 length of 74 mm (SD = 88 mm). The population E g lo sampled by the survey was strongly skewed to the right, with a modal size of 10 mm (figure 4). Few 5 fish were collected in the 100 to 140-mm size classes or in size classes above 350 mm. 0 0 50 100 150 200 250 500 350 400 450 500 The population structure in the bays was similar Length (mm) to that along the coast, being dominated by fish less than 100 mm SL (figure 5). However, the modal size was 10 mm along the coast and 20 mm in the bays. 30 I In addition, fish greater than 140 mm constituted I only a small portion of the population in the bays, Coast and no individuals of 240 mm or longer were taken. However, fish longer than 140 mm formed a signif- icant portion of the coastal population, which in- cluded many individuals longer than 240 mm. There were few fish between 100 and 140 mm SL in the coastal population. Distinct differences in the size-frequency distri- 5 bution of California halibut were apparent when "3 coastal sites were subdivided into semiprotected and 0 50 100 I50 200 250 300 350 400 450 500 exposed habitats (figure 6). In the exposed coastal Length (mm) habitat a strong mode was apparent at 10 mm; only Figure 5. Size distribution of California halibut (Paralichthys californicus) in three individuals were taken in the range of 20 to 140 bay and coastal habitats along the southern California coast, April-Septem- mm SL. In contrast, the size-frequency distribution ber 1989. in the semiprotected areas was similar to that for both coastal habitats combined, with many individ- uals between 20 and 140 mm SL; thus 10-mm and The densities of size groups varied from site to 150- to 300-mm fish constituted a smaller percent- site (figure 7). Settling and recently settled individ- age of the population. uals ((21 mm) were most dense in Anaheim Bay, followed by Hermosa Beach and Carlsbad. The densities of this size group were low at Long Beach and San Onofre, and the group was absent at Agua so IHedionda Lagoon. Larger age 0 fish (21-100 mm) 25 1 were more than twice as dense in Anaheim Bay as at Hermosa Beach, the area with the next greatest den- sity. Densities of this size group were low at Long Beach and Agua Hedionda Lagoon, and the group was virtually absent at Carlsbad and San Onofre. Older fish (>lo0 mm) were most dense at Hermosa Beach, followed by Carlsbad and Long Beach. The density of this size group was low in Agua Hedionda

0 50 100 150 200 250 300 350 40C 450 500 Lagoon and Anaheim Bay, and at San Onofre. In Length (mm) areas where it was present, the 21 to 100-mm group dominated. Figure 4. Overall size distribution of California halibut (Paralichthys californi- cus) in beam trawl surveys along the southern California coast, April-Sep- Changes in the population structure of California tember 1989. halibut with time differed between bay and coastal

89 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

so 250

Exposed 40- 2 200 "RI 0 0 - I 150 30 .-??z -c 0 0 e= & 20- g 5 100 n 5 10 - 5. 50

0 L !? Anaheim Bay Aguo Hsdlonda 0 50 100 150 200 250 300 350 400 450 501 Bay Sites Length (mm)

Semiprotected

- Hsrmoro Beach Long Beach Son Onofre Corlrbod Length (mm) Coastal Sites

Figure 6. Size distribution of California halibut (Paralichthys californicus) in Figure 7. Mean density (number of fishlha) of juvenile California halibut (Par- exposed and semiprotected coastal habitats along the southern California alichthys californicus) at sampling sites off southern California, April-Sep- coast, April-September 1989. tember 1989.

habitats (figures 8 and 9). In bays the primary mode September, settlement was highest at Anaheim Bay; consisted of 10-mm fish from April to June, and 20- from July to August it was highest at Hermosa to 40-mm fish from July to September (figure 8). Beach. In Anaheim Bay settlement was already high Fish of 50 mm SL or less were relatively abundant in in April and increased greatly from May to June. all months, and some larger fish were occasionally Settlement dropped dramatically in July, then re- captured. In August and September the percentage turned to relatively high levels again in August and of 50- to 100-mm fish increased. September. At Hermosa Beach settlement showed a In the coastal habitat, the 10-mm size class pre- gradual increase from April to August and fell to dominated from April to July, the 20-mm class in very low levels in September. Settlement was great- August, and the 50-mm class in September (figure est in May at Long Beach, and in July at Carlsbad 9). Although fish greater than 150 mm were present and San Onofre. in all months, individuals of 20 to 100 mm SL were California halibut settled into different locations rare until August and September, when they became at different temperatures (figure 11). In Anaheim abundant. However, this size group was found only Bay, settlement was high from 18.5" to 22.3"C, with at semiprotected coastal sites (Hermosa Beach and a peak at 21°C. Settlement dropped dramatically at Long Beach). 22.5"C (in July). However, at Hermosa Beach, set- The settlement of California halibut, as indicated tlement occurred between 15.5" and 19.5"C, with a by the monthly density of fish <21 mm, varied by peak at 16.8"C. At Carlsbad settling halibut were site and time (figure 10). From April to June and in found from 17" to 20.5"C, at Long Beach primarily

90 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

July 40 - n

Q p so

5 20 a

10

0 0 50 100 150 200 250 300 350 400 450 500 Length (mm) Length (mm)

50 50

August 40 40 Q I p 30 E, I 0 5 20 a I 10 10

n, 0 0 0 50 100 150 ZOO 250 300 350 400 450 500 0 50 100 I50 200 250 300 350 400 450 500 Length (mm) Length (mm)

June September 'O- n 40 -

Length (mm) Length (mm)

Figure 8. Monthly size distribution of California halibut (Paralichthys californicus) in bay habitats of southern California, 1989

at 17"C, and at San Onofre at 21.2"C. The regression DISCUSSION of density of settling individuals versus temperature Juvenile California halibut have been known for (y = 1.680~- 5.3; Y = 0.0096; r' = 0.009; d.E = some time to occur in bays of southern California. 34; figure 12) was not significant at p = 0.05. The Haaker (1975) found large numbers of juveniles in regression of the log (x + 1) density versus temper- Anaheim Bay and suggested that they remain there ature (y = -0.02~+ 1.436; Y = 0.088; r' = 0.007; until they reach about 200 mm, at which time they d.f. = 34) was also not significant. emigrate to the coast. Nets used in Haaker's study

91 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

60 60 , I

Aprll

Length (mm) Length (mm)

60

50 August 50 1 May I

10 -

Length (mm) Length (mm)

50 - June September 50 i

0 50 100 150 200 250 300 350 400 450 50 Length (mm) Length (mm)

Figure 9. Monthly size distribution of California halibut (Paralichthys califomicus) in coastal habitats off southern California, 1989.

included bag seines, commonsense seines, and small mesh size of 12.5 mm. Using similar gear, Plummer otter trawls with minimum mesh sizes of 3-6 mm et al. (1983) surveyed the nearshore zone off San (Klingbeil et al. 1975). Onofre and Oceanside at depths of 6-30 m. They No California halibut smaller than 150 mm were captured 1,580 California halibut, but fewer than found along the coast in 1972-73 in an extensive 2% were shorter than 100 mm SL. Fish smaller than otter trawl survey (342 samples) (M.J. Allen 1982). 100 mm are seasonally abundant in Elkhorn Slough, However, that study sampled depths from 10 to 200 Mugu Lagoon, Alamitos Bay, Anaheim Bay, New- m using small otter trawls with a minimum stretch- port Bay, Agua Hedionda Lagoon, and Mission Bay

92 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

County, although individuals have occasionally been noted there. Thejunior author of this study has observed small California halibut near the surf zone at Torrance. Using seines, beam trawls, and otter trawls with a minimum mesh size of 2 mm, L.G. Allen (1988a) found recently settled California halibut in pro- tected (Alamitos Bay) and semiprotected (Long Beach Harbor) habitats. With the same gear as that used in the present study, Kramer and Hunter (1987) found that California halibut <50 mm SL were abundant in Agua Hedionda Lagoon and Mission Bay in 1987; however, individuals of this size were Figure 10. Monthly density (number of fishlha) of recently settled (SL <21 mm) juvenile California halibut (Paralichthys californicus) at sampling sites rare along the open coast at depths of 6-14 m. In off southern California, 1989. 1988 densities were again relatively high in Agua Hedionda Lagoon and Mission Bay, and had in- creased in coastal areas, although they were low in

-* Curlrbad San Diego Bay (Kramer and Hunter 1988). Coastal p 200 - + sun Onolre settlement was highest at La Jolla and lowest at San m + Long BwCh Onofre (Kramer, in press). L.G. Allen et al. (in H * Hermora Baoch I 150- press) also found coastal settlement of California .-5- * Agua Hadiondo z% Anohcim Boy halibut in 1988. Among nearshore areas at depths of 5-10 m from Point Conception to San Mateo Point, ;6 IO0-- 5 age 0 California halibut (i.e., <80 mm) were most k 50- abundant in southeastern Santa Monica Bay (El Se- gundo and Malaga Cove) and Long Beach Harbor 0 (Belmont Shore); settlement was highest in July. 15 16 17 18 19 20 21 22 23 24 Temperature (“C) That study used the same type of beam trawl we used along the coast for this survey, with four 5-min Figure 11. Relationship of density (number of fish/ha) of recently settled (SL tows at a station. 121 mm) juvenile California halibut (Paralichthyscalifornicus) to temperature at sampling sites off southern California, April-September 1989. Kramer (in press) compared the selectivity of var- ious gears used in studies of small California halibut. Beach seines were found to be less effective than the Dsnrllv (Number per heclare) 200 1.0-m beam trawl in capturing juvenile halibut in x bays. The 1.0- and 1.6-m beam trawls were equally effective at capturing halibut <80 mm SL; however, the 1.6-m beam trawl was more effective than the 1.0-m beam trawl for collecting fish >80 mm. In the study reported here, the lack of large fish in bay catches is probably at least partly due to the de- creased effectiveness of the 1.0-m trawl used in bays, compared to the 1.6-m trawl used along the coast. Kramer (in press) also found that the 1.6-m beam trawl captured a greater proportion of halibut <200 15 16 17 18 19 20 21 22 23 24 Temperature (“C) mm SL, whereas the standard 7.6-m otter trawl used in southern California coastal surveys captured Figure 12. Relationship of density (number of fish/ha) of recently settled ju- venile (SL <21 mm) California halibut (Paralichthys californicus) to temper- more halibut >200 mm. This may partly account ature in surveys off southern California, April-September 1989. for the low catches of halibut of this size in otter trawl surveys (e.g., M.J. Allen 1982; Plummer et ai. (Haaker 1975; Plummer et al. 1983; Kramer and 1983). Hunter 1987; L.G. Allen 1988a; Kramer and Hunter The present study confirms the findings of L.G. 1988; Kramer, in press). Plummer et al. (1983) found Allen (1988a) that juvenile California halibut are no California halibut of <205 mm SL at depths less more abundant in bays and semiprotected coastal than 5 m (the surf zone) off northern San Diego locations than in exposed coastal locations. The

93 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

present study also substantiates findings of other that found in Alamitos Bay, which had the greatest studies (Kramer and Hunter 1987, 1988; L.G. Allen density from 1983 to 1985 (L. G. Allen 1988a). Com- et al., in press; Kramer, in press) indicating that hal- pared with sites sampled in 1987 (Kramer and ibut do settle in some places along the open coast in Hunter 1987), the density of halibut at Anaheim Bay some years. It also supports L.G. Allen et al. (in in 1989 was about 50% of that found in Agua He- press), who found the greatest coastal settlement in dionda Lagoon (the area of greatest settlement) and southeastern Santa Monica Bay and Long Beach was similar to that of Mission Bay (Kramer and Harbor. It also confirms studies (Kramer and Hunter 1987). Settlement in Mission Bay and Agua Hunter 1987; L.G. Allen 1988a; Kramer and Hunter Hedionda Lagoon dropped in 1988. In 1989 no re- 1988; Kramer, in press) indicating that successful cently settled halibut were taken at Agua Hedionda settlement along the coast is low in the San Onofre- Lagoon, although larger age 0 fish (21-100 mm SL) Carlsbad region. were present at low densities. Thus settlement of L.G. Allen (1988a) found that the Los Angeles- California halibut to bays varies internannually, and Long Beach Harbor provides a nursery for Califor- the importance of a given bay as a nursery changes nia halibut. This finding is not entirely unexpected, from year to year. because breakwaters protect this area from swells In this study the density of recently settled Cali- and large waves from the west. However, the im- fornia halibut along the coast at Hermosa Beach was portance of southeastern Santa Monica Bay as a about 50% of that found at Anaheim Bay. The den- nursery is less obvious. Southeastern Santa Monica sity at Hermosa Beach in 1989 was about 100 times Bay is an open coast that is directly exposed to west- that found along the open coast off San Diego ern swells (Maloney and Chan 1974). However, in County in 1987 (Kramer and Hunter 1987) and 2-3 mid to late summer the area is protected from the times greater than that found there in 1988 (Kramer predominant south and southwestern swells by the and Hunter 1988). It was slightly less than that found Palos Verdes Peninsula (Maloney and Chan 1974). at Malaga Cove in 1988 (L.G. Allen et al., in press). Juvenile California halibut have previously been Kramer (in press) noted that although settlement observed in this area, but were not documented un- might be high along coastal San Diego County early til recently. As noted above, small California halibut in the year, few halibut of 40-60 mm were taken have been observed at Torrance in the past, and in later. Possible reasons for their absence include mor- some years juveniles have been found in the seawater tality, dispersal to deep water, and dispersal to bays. cooling system of the Redondo Generating Station Kramer surmised that dispersal to bays was most (M.D. Curtis, MBC Applied Environmental Sci- likely because (1) there is no evidence of dispersal to ences, Costa Mesa, Calif, pers. comm.). In 1988 deep water, and (2) more fish of 21-100 mm were settlement was particularly high at Malaga Cove and found in bays than fish <21 mm, indicating that El Segundo (L.G. Allen et al., in press); these sites settlement must be occurring elsewhere. However, were north and south, respectively, of the Hermosa we believe that mortality may be an important factor Beach site used in this study. We found settling hal- in exposed coastal areas where there are no suitable ibut at Hermosa Beach from April to September. bay habitats. However, survival (or residency) of these fish only The relatively high survival (or residency) of age occurred there from August to September, when the 0 fish near Hermosa Beach in 1989 may be related to Palos Verdes Peninsula provided the greatest protec- the proximity of King Harbor, which may provide tion from southern swells. needed protection, especially in times of greater At least two semiprotected coastal areas of south- exposure. ern California - southeastern Santa Monica Bay and L. G. Allen (in press) found that log-transformed Los Angeles-Long Beach Harbors -seem to pro- abundance of settling halibut along the coast was vide suitable nursery grounds for California halibut significantly and positively correlated with temper- in some years, particularly late in the summer. The ature. That study concluded that temperature has a coast north of the La Jolla Peninsula may be another significant influence on the settlement and subse- suitable area that is somewhat protected from swells quent distribution of age 0 halibut. But the present from the south. Kramer (in press) found greater hal- survey did not show any significant correlation with ibut settlement there (at Torrey Pines) than at San temperature. This lack of correlation may partly re- Onofre. flect the more heterogeneous study area that includes The site with the greatest density of recently set- both bay and coastal sites. However, in the present tled California halibut in this study was Anaheim study settlement was often greatest at intermediate Bay. The density there in 1989 was about 25% of temperatures at a given site (e.g., Anaheim Bay,

94 ALLEN AND HERBINSON: SETTLEMENTOF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

Hermosa Beach; figure 11). Other physical or bio- (lab, field, graphics); Janalyn Sanders (data analysis, logical factors are apparently more important than graphics); and Bill Stockton (data analysis, graph- temperature in determining settlement at some sites. ics). The following members of the MBC staff as- Variations in pH are generally directly related to sisted in the lab: Pat Frost, Scott Harris, Jenny dissolved oxygen (DO) levels (Parsons and Taka- Higuera, Diana Budris, Julie Smith, Steve Burke, hashi 1973). When DO levels are low, pH values are Lisa Kasper, Leslie Keyt, and Tiffany Woodworth. generally low; when DO values are high, pH levels Other MBC staff members helped in the following are high. Both DO levels and pH values are high areas: graphics (Larry Jones); word processing (Gini when photosynthetic rates are high, and both are Whitt, Phyllis Barton, and Madine Johnson); report low when respiration predominates. The high pH editing (Tom Kauwling); and Spanish editing (Ka- observed along the coast in August and September thy Mitchell and Joan Compton). and in the bays in May, July, and August may reflect We would also like to thank Sharon Kramer (now high photosynthesitic activities. There does not ap- a member of the MBC staff) and John Hunter of the pear to be any obvious relationship of settlement Southwest Fisheries Center, National Marine Fish- to pH. Settlement was highest at Anaheim Bay dur- eries Service, NOAA, La Jolla, California, for lend- ing the period of lowest pH but was highest at Her- ing their beam trawls and meter wheels for use in mosa Beach during a period of high pH (table 1; this project. We appreciate the help of Larry Nufer figure 10). of Conshelf Services Inc., who was skipper of the R/V Westwind. We also appreciate the technical assis- CONCLUSIONS tance of Marty Golden, who volunteered to help California halibut settled from the plankton to the with the project on his own time. bottom in shallow coastal areas off southern Cali- fornia in 1989 in varying densities depending on habitat type. LITERATURE CITED Allen, L. G. 1988a. Recruitment, distribution, and feeding habits of Transforming larvae were found throughout the young-of-the-year California halibut (Paralichthys califomicus) in the study area, but the abundance (survival and resi- vicinity of Alamitos Bay-Long Beach Harbor, California, 1983- dency) of small juveniles was highest in the bays, 1985. S. Calif. Acad. Sci., Bull. 87(1):19-30. -. 1988b. OREHP YOY white seabass trawl survey, 1988, Con- intermediate in semiprotected coastal areas, and tract #FG7500, quarterly report for April-June 1988. Prepared for lowest in exposed coastal areas. Calif. Dep. Fish and Game, Long Beach. Calif. State Univ.. North- Benthic settlement diminished in August and ridge, Calif., 4 pp. Allen, L. G., R. E. Jensen, and J. R. Sears. In press. Open coast recruit- September, when larger young-of-the-year became ment and distribution of young-of-the-year California halibut (Par- more abundant; these larger fish did not appear at alichthys californictrs) along the southern California coast between Pt. exposed coastal locations. Conception and San Mateo Pt.,June-October, 1988. Calif. Dep. Fish Game, Fish. Bull. The results ofthis study substantiate the results of Allen, M. J. 1982. Functional structure ofsoft-bottom fish communities other recent studies indicating that southeastern of the southern California shelf. Ph.D. dissertation, Univ. Calif., San Santa Monica Bay and Los Angeles-Long Beach Diego, La Jolla, Calif., 577 pp. (available from Univ. Microfilms Internatl., Ann Arbor, MI. Ref. 8300991). Harbors are semiprotected coastal areas that are used CDFG (California Department of Fish and Game). 1989. Review of as nursery areas by California halibut. This study some California fisheries for 1988. Calif. Coop. Oceanic Fish, Invest. does not, however, substantiate a correlation be- Rep. 30:7-17. Haaker, l? L. 1975. The biology of the California halibut, Paralichthys tween temperature and settlement. caliJornicns (Ayres) in Anaheim Bay. In The marine resources of Ana- heim Bay, E. D. Lane and C. W. Hill, eds. Calif. Dep. Fish Game, Fish Bull. 165:137-151. ACKNOWLEDGMENTS Klingbeil, R. A,, R. D. Sandell, and A. W. Wells. 1975. An annotated This project was funded by Southern California checklist of the elasmobranchs and teleosts of Anaheim Bay. In The Edison Company as part of its Natural Resources marine resources of Anaheim Bay, E. D. Lane and C. W. Hill, eds. Calif. Dep. Fish Game, Fish Bull. 165:79-115. Management Research Program. We would like to Kramer. S. H. In press. Distribution and abundance ofjuvenile Califor- thank Chuck Mitchell of MBC Applied Environ- nia halibut (Paralichthys caliJornicur) off San Diego County, California. mental Sciences for providing the personnel, facili- Calif. Dep. Fish Game, Fish Bull. Kramer, S. H., and J. R. Hunter. 1987. Southern California wetland/ ties, and technical assistance to conduct this study. shallow water habitat investigation, annual report for the fiscal year The following members of the MBC staff helped in 1987. U.S. Dep. Commerce, NOAA, NMFS, SWFC, La Jolla, Calif., a variety of technical areas: Bob Moore (field chief, 12 pp. -. 1988. Southern California wetland/shallow water habitat in- lab, data analysis); Tina Moskowitz (lab, field); vestigation, annual report for the fiscal year 1988. U.S. Dep. Com- Mark Burke (field, lab, data analysis); Don Johnston merce, NOAA, NMFS, SWFC, La Jolla, Calif., 15 pp.

95 ALLEN AND HERBINSON: SETTLEMENT OF CALIFORNIA HALIBUT, 1989 CalCOFl Rep., Vol. 31,1990

Maloney, N. J., and K.-M. Chan. 1974. Physical oceanography. In A NMFS (National Marine Fisheries Service). 1985. Marine recreational summary of knowledge of the southern California coastal zone and fishery statistics survey, Pacific Coast, 1983-1984. U.S. Dep. Com- offshore areas. Vol. I, Physical environment. M. D. Dailey, B. Hill, merce, NOAA, NMFS, Washington, D.C., 189 pp. and N. Lansing, eds. Prepared for U.S. Dep. Interior, Bur. Land Parsons, T. R., and M. Takahashi. 1973. Biological oceanographic pro- Mgmt. by So. Calif. Ocean Studies Consortium, Calif. State Univ. cesses. New York Pergamon Press, 186 pp. & Coll., Dominguez Hills, Calif., Contract no. 08550-CT4-1, pp. 3- Plummer, K. M., E. E. DeMartini, and D. A. Roberts. 1983. The 1 to 3-65. feeding habits and distribution ofjuvenile-small adult California hal- MBC Applied Environmental Sciences. 1990. Distribution ofjuvenile ibut (Paralichthys calijornicus) in coastal waters off northern San Diego California halibut (Puralichthys cul~o~nicus)and other fishes in bay and County. Calif. Coop. Oceanic Fish. Invest. Rep. 24:194-201. coastal habitats of Los Angeles, Orange, and San Diego counties, in 1989. Prepared for S. Calif. Edison Co., Rosemead, Calif. MBC Applied Environ. Sci., Costa Mesa, Calif. 90-RD-09.74 pp.

96 BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

GENETIC STRUCTURE OF WHITE SEABASS POPULATIONS FROM THE SOUTHERN CALIFORNIA BIGHT REGION: APPLICATIONS TO HATCHERY ENHANCEMENT

DEVIN M. BARTLEY DONALD B. KENT Department ofAiiimal Science Sea World Research Institute University of California 1700 South Shores Drive Davis, California 95616 San Diego, California 92109

ABSTRACT de la variabilidad genCtica del robalo blanco fueron Although some fisheries biologists have expressed evaluadas con “starch gel” electroforesis horizontal concern over the practicality of hatchery enhance- en nueve Areas costeras del sur de California (South- ment of marine fisheries, hatchery technology has ern California Bight). El proposit0 de este estudio advanced significantly during the 1980s. Marine es el de describir la estructura genttica del robalo hatchery enhancement programs are now address- blanco y utilizar esta informacion en el mejora- ing ecology and genetics as well as animal husband- miento de 10s criaderos. La heterocigosidad prome- ry. To describe the genetic structure of white dio observada en cada muestra fue de 0.024 a 0.064. seabass and apply the information to hatchery en- Los indices de identidad genttica entre muestras fu- hancement, we used starch-gel electrophoresis to as- eron mayores de 0.99. Dado que el 3% de la hetero- sess the level and distribution of genetic variability cigosidad total se debi6 a las diferencias entre las of seabass from nine areas in the Southern California muestras y dado que el nfimero estimado de indivi- Bight. Average heterozygosity per sample estimates duos migratorios con intercambio genttico entre las ranged from 0.024 to 0.064. Indices of genetic iden- muestras fue aproximadamente de 9, creemos que tity between samples were greater than 0.99. Be- existe muy poca subdivision en la poblacion del Area cause only 3% of the total gene diversity was due to estudiada. La diversidad genktica en seis muestras intersample differences, and because an estimate of de robalo de criadero fue 15% menor que la obser- number of migrants exchanging genes among sam- vada en la poblacion silvestre. Esto se debe proba- ples was approximately 9 per generation, we believe blemente a1 numero reducido de reproductores little population subdivision exists within the study utilizados en 10s desoves en masa. A pesar de que la area. Genetic diversity in six hatchery samples of estructura genCtica de la progenie obtenida en cada white seabass was 15% lower than that found in the desove en masa pueda diferir de las muestras de po- wild, possibly because small numbers of brood blaciones silvestres, desoves consecutivos incremen- stock contribute to mass spawns. Although the ge- taron la variabilidad genktica cuado se considera la netic structure of progeny from a single mass spawn production total del criadero. Por lo tanto, 10s de- may differ from wild samples, successive mass soves consecutivos durante el period0 del desove del spawns increased the genetic variability of the entire robalo blanco parece ser un mCtodo efectivo para hatchery product. Therefore, continuous mass mantener la diversidad genCtica en 10s criaderos. spawning of white seabass over the course of the spawning season appears to be an effective means of INTRODUCTION preserving genetic diversity. At the 1988 meeting of the California Cooperative Oceanic Fisheries Investigations, some attendees ex- RESUMEN pressed concern about the practicality of hatchery A pesar de las dudas expresadas por 10s biologos enhancement of marine fisheries. In light of past pesqueros con respecto a la utilidad de las piscifac- attempts at enhancement, much of this concern is torias para el mejoramiento de las pesquerias mari- justified. Many early hatchery enhancement pro- nas, la tecnologia de estos criaderos ha mostrado grams had no demonstrable effect on the target pop- avances significativos en la iiltima dCcada. Hoy dia ulation (see references in MacCall 1989). In the case 10s programas de mejoramiento dirigen su atencidn of Pacific salmon, many hatchery programs were hacia la ecologia y la genttica, asi como tambiCn developed to mitigate habitat degradation resulting hacia la cria de ios animales. El nivel y la distribucibn from water development projects (Netboy 1974). Clearly, failing to evaluate a hatchery’s contribution to the target population, and using hatcheries to jus- ~~ ~~ [Manuscript received February 1,1990 ] tify habitat destruction are unacceptable.

97 BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

However, hatchery enhancement programs are sected at Sea World Reseach Institute, and tissue now addressing ecological and genetic concerns as samples were frozen and stored at - 70" C. well as animal husbandry, and evaluation programs We also collected juvenile white seabass from six are being implemented (Rutledge 1989). Biochemi- separate mass spawns at Sea World Research Insti- cal genetics has become one important tool in the tute (SWRI): one sample in 1985 (N = 21); one sam- study of fishery biology and (Ryman ple in 1986 (N = 72); and four samples in 1988 (N's and Utter 1987). Applying biochemical genetic = 30, 26, 27, and 36). These hatchery samples were techniques to hatchery enhancement will help the designated SWRI-1 through SWRI-6. All six hatch- hatcheries produce viable fish by preserving genetic ery samples of white seabass descended from ap- diversity (Kincaid 1983; Allendorf and Ryman proximately 20 brood stock maintained in a 3.0 x 1987). At the same time, the use of genetic markers 6.0 x 1.2-m pool (19,000 liters) with recirculating will increase fishery scientists' ability to evaluate seawater. Adult white seabass were induced to hatchery contribution to wild populations (Murphy spawn by photoperiod and temperature manipula- et al. 1983; Seeb et al. 1986). tion. Eggs were collected by siphon, and larvae The objective of the research reported here is to hatched after 2-3 days. White seabass progeny were apply biochemical and population genetic tech- fed a combination of rotifers, Artemia, and commer- niques to hatchery enhancement efforts on white cially available trout feed until they were approxi- seabass, Atractoscion nobilis (Scianidae), in the South- mately 9 cm long, at which point they were frozen ern California Bight region. Numbers of this species whole and stored at -70°C. Muscle and liver tis- in the Southern California Bight have been declin- sues were dissected and stored at -70°C before ing, and the species has almost completely disap- electrophoresis. peared from the central California waters that historically supported the center of the commercial Electrophoresis fishery (Skogsberg 1939; Vojkovich and Reed 1983). Horizontal starch-gel electrophoresis followed We use allozyme data from horizontal starch-gel standard procedures (Aebersold et al. 1987). En- electrophoresis of soluble proteins to describe the zyme systems, tissue distribution, and number of level and distribution of genetic diversity in natural loci are presented in table 1. Gels were made with and cultured populations of white seabass. Al- 12% hydrolyzed potato starch (Sigma Chemical though these techniques have been used to describe Co.) combined with one of the following buffer sys- the population genetic structure of Gulf of Mexico tems: AC, an amine citrate buffer from Clayton and Scianidae (Ramsey and Wakeman 1987), genetic Tretiak (1972) adjusted to pH 7.0; R, a discontin- studies of marine fish in the Southern California uous buffer system (pH 8.0) described by Ridgeway Bight have, until now, neglected white seabass et al. (1970); and MF, a boric acid-Tris system (pH (Waples 1987). 8.4) from Markert and Faulhaber (1965). For histochemical staining procedures we fol- METHODS lowed Shaw and Prasad (1970) and Harris and Hop- kinson (1976). For nomenclature we followed Collections and Samples Allendorf and Utter (1979); in this system, the most We collected 13 samples of juvenile and adult common allele at a locus is designated the 100 allele, white seabass from the Southern California Bight and variant alleles are assigned numeric values based region using gill nets of approximatley 3.8-, 7.6-, on their anodal migration distance relative to the 100 and 8.9-cm mesh. Specific locations and dates of allele. Enzyme abbreviations are in uppercase letters; collections were: Point Loma 1988 (Sample size [N] alleles are designated by the italicized enzyme abbre- = 40); Point Loma 1987 (N = 50); Mission Bay, San viation followed by locus number with allelic mo- Diego 1988 (N= 36); La Jolla 1988 (N = 23); La Jolla bility in parentheses. 1987 (N = 90); Encinitas 1988 (N = 5); Encinitas 1987 (N = 100); San Onofre 1988 (N = 35); San Data Analysis Onofre 1987 (N = 16); Seal Rock, San Clemente We assessed genetic variability of each fish sample 1988 (N = 51); San Mateo Point, San Clemente 1988 by calculating the frequencies of alleles at each locus, (N = 34); Dana Point 1988 (N = 17); and Laguna the percentage of polymorphic loci (P),and average Beach 1988 (N = 13). Muscle and liver tissue were heterozygosity (H) (Nei 1973). A locus was consid- dissected from the fish in the field and frozen in ered polymorphic if we observed one variant allele. liquid nitrogen. Fish from Mission Bay were dis- We calculated genetic identities (I)for each pair of

98 BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

TABLE 1 Enzyme Systems and Isozyme Loci Analyzed from Liver (L) and Muscle (M) Tissue of White Seabass Number Enzyme system Abbreviation E.C. number of loci Tissue Buffer* Acid phosphatase ACP 3.1.3.2 2 MJ- MF Aconitate hydratase Ah 4.2.1.3 2 L AC Adenylate kinase Ak 2.7.4.3 1 M AC Alcohol dehydrogenase Adli 1.1.1.1 1 L AC,MF Aspartate amino transferase Aat 2.6.1.1 2 M,L AC,R Creatine kinase Ck 2.7.3.2 1 M R Esterase Est 3.1.1.1 2 M, L R Glyceraldehyde-3-phosphate dehydrogenase Gapdh 1.2.1.12 2 M AC Glycerol-3-phosphate dehydrogenase GPd 1.1.1.8 -3 M AC Glucose phosphate isomerase Cpi 5.3.1.9 3 M R Iditol dehydrogenase Iddh 1.1.1.14 1 L R Isocitrate dehydrogenase Idh 1.1.1.42 2 M,L AC Lactate dehydrogenase Ldh 1.1.1.27 1 M R Malate dehydrogenase Mdh 1.1.1.37 1 M,L AC Malic enzyme Me 1.1.1.40 1 MJ- MF Mannose phosphate isomerase Mp i 5.3.1.8 1 M,L R,MF Phosphogluconate dehydrogenase 6 Pdg 1.1.1.44 1 M, L AC Phosphoglucomutase Pgm 5.4.2.2 1 M,L AC Superoxide dismutase Sod 1.15.1.1 1 M, L MF, R Triosphosphate isomerase TP1 5.3.1.1 -3 M MF Peptidase Glycyl leucine DPeP 3.4.13.11 2 M R Leucyl glycyl glycine TaPeP 3.4.11.4 1 M R ~______*Described in text

samples (Nei 1978) and constructed a dendrogram generation. Equation 1was solved for Nrn by setting from estimates of I with the unweighted pair-group FST equal to the relative gene diversity (Gs,) esti- method (UPGMA; Sneath and Sokal 1973) to ex- mate (Nei 1977). Equation 1 will provide an estimate amine the relative similarities among populations. of the number of migrant fish exchanging genes We partitioned total gene diversity (H,) of wild among samples per generation under the assump- and hatchery samples to estimate within-sample tions of selective neutrality of alleles and Wright’s (23,)and between-sample (D,,) components and (1943) island model of migration. Slatkin and Barton relative gene diversity ( G,,) (Nei 1973; Chakra- (1989) discussed the sensitivity of equation 1 to var- borty and Leimar 1987). To ascertain significant sub- ious methods of estimating Fsr, some selection, and population structure within the study area, we used population structure, and found it to be fairly a chi-square test to determine if Wright’s FST (1943) robust. was significantly different from zero, as described by Waples (1987). We used G,, to approximate F,, RESULTS (Nei 1977; Slatkin and Barton 1989). Tests of inde- pendence between allelic frequency and location Allozyme Variation of Wild Samples were performed by log likelihood G test (Sokal and We detected allozyme variation in 19 of 33 loci Rohlf 1981). G tests were also performed on allelic (table 2). The distribution of alleles in wild samples frequencies and mass spawning samples to deter- of white seabass was generally homogeneous mine if significant genetic differences existed among throughout the study area, except for rare alleles in the six hatchery samples. Quantitative estimates of specific locations. For example, we observed the gene flow were calculated from Wright’s (1943) fix- Gpd-2 (- 233) allele only in the 1987 sample from San ation index Onofre. In addition, IDH, ME, and MPI poly- morphisms were observed in 1988 and not in 1987 FST = 1/(4Nm + 1) (1) samples. The distribution of rare alleles did not fol- low an obvious pattern. Heterozygosity estimates where Nm is the average number of migrants per ranged from 0.033 in the Encinitas 1988 sample to

99 BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

TABLE 2 Allelic Frequencies at 19 Polymorphic Gene Loci from 13 Wild and 6 Hatchery Samples of White Seabass - Pt. Loma Pt.Loma Mission Bav La Tolla La Jolla Encinitas Encinitas San Onofre San Onofre 1988 1987 1988 1988 1987 1988 1987 1988 1987 ~~ ~~ ~ ~~~ ~~ ~ ~~~ ~~ ~ ~~ ~ Aat-1 (100) 0.975 0.906 0.930 1.000 0.909 0 900 0.870 0.886 0.860 (115) 0.025 0.021 0.028 0.025 0 100 0.019 0.071 0.031 (50) 0.073 0.042 0.066 0.111 0.043 0.109 N* 40 48 36 23 90 5 79 35 16 Aat-2 (100) 0.872 0.995 0.944 0.935 0.945 1 000 0.992 0.914 1.000 (125) 0.103 0.005 0.028 0.043 0.049 0.004 0.086 (65) 0.026 0.028 0.022 0.006 0.004 N 39 50 36 23 86 5 100 35 16 Adh (-100) 0.462 0.443 0.486 0.304 0.543 0 500 0.438 0.443 0.385 ( - 200) 0.487 0.534 0.514 0.652 0.429 0 500 0.531 0.557 0.615 ( - 300) 0.051 0.023 0.043 0.028 0.031 N 39 44 36 23 35 5 81 35 13 Est-4 (100) 0.936 0.896 0.861 0.978 0.927 1 000 0.950 0.914 0.962 (89) 0.010 0.056 0.028 0.025 0.038 (115) 0.064 0.094 0.083 0.022 0.045 0.025 0.086 N 39 48 36 23 89 5 40 35 13 Est-5 (100) 0.938 1.000 0.875 0.957 0.925 1 000 1.000 0.900 0.937 (103) 0.063 0.083 0.043 0.050 0.071 (95) 0.042 0.025 0.029 0.063 N 40 20 36 23 40 5 40 35 16 Gpd-2 (- 100) 1.000 1.000 1.000 1.000 1,000 1 000 1.000 1.000 0.967 (- 133) 0.033 N 40 50 36 23 90 5 100 35 16 Gpr-l (100) 1.000 1,000 1,000 1.000 1.000 1 000 1.000 1.000 1.000 (- 140) N 40 50 36 23 90 5 80 35 16 Gpi-2 (100) 0.975 1.000 0.958 1.000 0.994 1 000 1,000 1.000 1.000 (85) 0.025 0.006 (122) 0.042 N 40 50 36 23 90 5 80 35 16 Gpi-3 (100) 1.000 1,000 1.000 1.000 0.994 1 000 0.987 1.000 1.000 (78) 0.006 0.013 (112) N 40 50 36 23 90 5 80 35 16 Iddh-1 (100) 0.988 0.970 1.00 0.957 0.970 1 000 0.990 0.943 1,000 ( - 400) 0.043 0.030 0.005 0.043 (700) 0.012 0.030 0.005 0.014 N 40 50 36 23 50 5 100 35 16 Idh-1 (100) 1.000 1.000 0.986 0.978 1.000 1 000 1.000 0.986 1.000 (82) 0.014 0.022 0.014 N 40 50 36 23 90 5 100 35 16 Idh-2 (100) 0.975 1,000 0.986 1.000 1.000 1 000 1.000 0.986 1,000 (119) 0.025 0.014 0.014 (62) N 40 50 36 23 90 5 100 35 16 Ldh-3 (100) 0.950 1,000 0.972 0.978 0.983 1 000 0.980 0.986 0.934 (40) 0.025 0.014 0.017 0.015 0.014 0.033 (150) 0.025 0.014 0.022 0.005 0.033 N 40 50 36 23 88 5 100 35 16 Me1 (100) 0.975 1.000 1.000 0.935 1.000 1 000 1.000 1,000 1.000 (116) 0.025 0.065 N 40 50 36 23 90 5 100 35 16 Mpi (100) 0.949 1.000 0.917 0.978 1.000 1 000 1.000 0.971 1.000 (110) 0.051 0.083 0.022 0.029 N 39 50 36 23 90 5 100 35 16 6Pgd (100) 1.000 0.980 1.000 1.000 1.000 1 000 0.995 1.000 1.000 (95) 0.020 0.005 N 40 50 36 23 90 5 90 35 16 Pgm I ( - 100) 0.224 0.410 0.403 0.239 0.250 0 300 0.237 0.371 0.447 (- 167) 0.750 0.590 0.555 0.761 0.750 0 700 0.750 0.586 0.553 ( - 200) 0.026 0.042 0.013 0.043 N 38 50 36 23 10 5 40 35 16 Sod (100) 0.988 1.000 0.986 1.000 0 989 1 000 1,000 1.000 1.000 ( 109) 0.012 0.014 0.011 N 40 48 36 23 88 5 100 35 16 Ah-2 (100) 1.000 1,000 1,000 1.000 1.000 1 000 0.990 1.000 1.000 (98) 0.010 N 40 50 36 5 90 5 99 35 16 H 0.056 0.045 0.064 0.043 0.048 0 033 0.041 0.060 0.048 P 0.39 0.21 0.36 0.30 0.33 0 09 0.30 0.33 0.21

~~ *Sample size (continued on nextpage) H = average heterozygosity; P = proportion polymorphic loci BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

TABLE 2 (continued)

Seal Rock San Mateo Pt. Dana Pt Laguna Beach SWRI-I SWRI-2 SWRI-3 SWRI-4 SWRI-5 SWRI-6 1988 1988 1988 1988 1985 1986 1988 1988 1988 1988 ___ ~ Aut-I (100) 0.800 0.783 0.941 1.000 0.658 1.000 0.917 0.923 0.833 0.853 (115) 0.030 0.152 0.059 0.250 0.033 0.019 0.015 (50) 0.170 0.065 0.092 0.050 0.077 0.148 0.132 N* 50 23 17 13 21 72 30 26 27 34 Aut-2 (100) 0.908 0.912 0.969 0.923 1.000 1.000 0.967 1.000 0.963 0.686 (125) 0.071 0.059 0.031 0.077 0.037 0.300 (65) 0.020 0.029 0.033 0.014 N 49 34 16 13 21 35 30 26 27 35 Adh (-100) 0.357 0.411 0.375 0.538 0.833 0.910 0.517 0.923 1.OOO 0.588 ( - 200) 0.633 0.574 0.594 0.462 0.167 0.090 0.483 0.077 0.412 ( - 300) 0.010 0.015 0.031 N 51 34 16 13 21 72 30 26 27 34 Est-4 (100) 0.930 0.941 0.938 0.962 1,000 1.000 1.000 1.000 1.000 0.958 (89) 0.020 0.029 (115) 0.050 0.029 0.062 0.038 0.042 N 50 34 16 13 21 72 30 26 27 36 Est-5 (100) 0.940 0.958 0.912 1.000 1.000 1.000 1,000 1.000 1.000 0.771 (103) 0.040 0.021 0.088 (95) 0.020 0.021 0.229 N 49 24 17 13 21 72 30 26 27 35 Gpd-2 (- 100) 1,000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 I.000 (- 133) N 51 34 17 13 21 72 30 26 27 35 Gpi-I (100) 1.000 1.000 1,000 1.000 1.000 0.708 1.000 1.000 1.000 1,000 (- 140) 0.292 N 51 34 17 13 21 72 30 26 27 36 Gpi-2 (100) 1.000 1.000 1.000 1.000 1.000 1.000 1,000 1.000 1.000 1.000 (85) (122) N 51 34 17 13 21 72 30 26 27 35 Gpi-3 (100) 1,000 0.971 0.971 1.000 1.000 1.000 1.000 1.000 1.000 1.000 (78) 0.029 (112) 0.029 N 51 34 17 13 21 72 30 26 27 35 Iddh-2 (100) 0.979 0.985 1,000 0.924 1.000 1.000 1.000 1.000 1.OOO 1.000 ( - 400) 0.021 0.015 0.038 (700) 0.038 N 48 34 16 13 21 72 30 26 27 35 Idh-I (100) 1.000 0.957 1.000 1.000 1.000 1.000 0.983 1.000 0.870 1.000 (82) 0.043 0.017 0.130 N 51 23 17 13 21 72 30 26 27 36 Idh-2 (100) 1,000 0.979 1.000 1.000 1.000 1.000 1.000 1.000 0.944 1.000 (119) (62) 0.021 0.056 N 51 24 17 13 21 72 30 26 27 36 Ldh-3 (100) 0.990 1,000 0.941 1,000 1.000 0.972 1.000 1.000 1.000 1.000 (40) 0.010 0.029 0.028 (150) 0.029 N 51 34 17 13 21 72 30 26 27 36 Me-I (100) 1.000 1.000 1.000 1.000 1.000 1.000 1,000 1.000 1.000 1,000 (116) N 51 34 17 13 21 72 30 26 27 36 Mpi (100) 0.957 0.850 0.969 1.000 1.000 0.965 1.000 1.000 1.000 1.000 (110) 0.043 0.150 0.031 0.035 N 47 30 16 13 21 72 30 26 27 36 6Pgd (100) 0.990 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 (95) 0.010 N 50 34 17 13 21 72 30 26 27 36 Pgm-f (-100) 0.352 0.221 0.406 0.192 0.857 0.743 0.283 0.462 0.442 0.750 (- 167) 0.637 0.735 0.594 0.769 0.143 0.257 0.667 0.538 0.558 0.250 ( - 200) 0.011 0.044 0.038 0.050 N 44 34 16 13 21 72 30 26 26 32 Sod (100) 1.000 0.985 0.971 1.000 1.000 1.000 1.000 1.000 1,000 1,000 (109) 0.015 0.029 N 38 34 17 13 21 72 30 26 27 36 Ah-2 (100) 1.000 1.000 1.000 0.923 1.000 1.000 1.000 1.000 1.000 1.000 (98) 0.077 N 49 29 11 13 21 72 30 26 27 36 H 0.056 0.064 0.052 0.042 0.033 0.03 0.037 0.024 0.036 0.060 P 0.30 0.36 0.30 0.18 0.09 0.15 0.15 0.09 0.15 0.18 BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

0.064 in the San Mateo Point and the Mission Bay method of assessing genetic differences between samples. The proportion of polymorphic loci was subpopulations. We used G,, to approximate F,, lowest in the Encinitas 1988 sample and highest in (Nei 1977) and found highly significant F,, values the Point Lorna 1988 sample. However, because only in both the 1987 (F,, = 0.0304; d.f = 72; XZ = five fish were collected from Encinitas in 1988, the 247.953) and 1988 (FsT = 0.0259; d.f = 240; x2 = estimates of genetic variability may be inaccurate. 399) samples. However, tests of independence of al- Cluster analysis based on Nei’s genetic identity lele frequency and location on the 1987 and 1988 data estimates failed to reveal any geographic structure sets revealed significant association of alleles and lo- to the genetic variation in white seabass (figure 1). cation only for Aut-1 in 1988 (G = 29.391; d.f = 16). Genetic identity values between wild samples were To determine if significant allele frequency differ- all greater than 0.99. ences existed between samples taken at different To determine if population substructuring was years from the same location, tests of independence occuring in natural populations of white seabass, we of allele frequencies and sample year were performed examined gene diversity analysis quantifying the on collections from Point Loma, La Jolla, Encinitas, amount of genetic variation between (D,,) and and San Onofre. Although the summary indices of within (H,) samples. For natural populations of genetic variation changed between years (H and P, white seabass sampled in 1987 and 1988, approxi- table 2), only the Aut-3 locus from the Point Loma mately 97% of the variation was derived from samples displayed significant temporal heterogene- within-sample variability (DsT = 0.0017; H, = ity (G = 8.518; d.f = 1). 0.0535; H, = 0.0552 for 1987 samples, and D,, = Estimates of gene exchange were high among 0.0013; H, = 0.0487; H, = 0.0500 for 1988 sam- wild samples. We estimated the level of gene flow ples) and 3% was due to between-sample differences among wild samples to be 8.0 individuals per (G,, = 0.304 for 1987 and 0.0259 for 1988). generation in 1987, and 9.4 in 1988 (equation 1). In spite of this low level of intersample variability, we did find statistical evidence of differences in ge- Allozyme Variability in Hatchery Samples netic variability. Wright’s (1943) F,, statistic mea- We observed eleven polymorphic loci in six hatch- sures the reduction in heterozygosity observed in a ery samples of white seabass. However, individual population due to population subdivision and is one samples had as few as three, and no more than six polymorphic loci (table 2). Heterozygosity levels Point Lorna 1988 ranged from 0.024 to 0.060 and were comparable to Le Jolla 1987 levels in natural samples. Encinitas 1988 Although white seabass progeny all came from a SWRI-3 1988 common brood stock, differences in allelic variabil- Laguna Beach 1988 ity were apparent. For example, SWRI-2 was the Encinitas 1987 only hatchery sample in which we observed the Ldh- La Jolla 1988 3(40),Gpi-l( - 140),and Mpi-(110)alleles. However, Point Lorna 1988 no Aut-1 or Aut-2 polymorphisms were seen in this nission BOY 1988 sample. SWRI-5 was fixed for the common Adh al- Sen Onofre 1988 lele, whereas other hatchery samples were variable san Onofre 1987 at this locus. The Est-5 locus was polymorphic only Dana Point 1988 in SWRI-6 sample. Seal ROCk 1988 Gene diversity analysis of hatchery samples indi- San nateo Pt 1988 cated 84% of the total heterozygosity was due to SWRI-6 1988 differences within samples, whereas 16% (G,, = SWRI-4 1988 0.157) originated from between-sample differences, I r SWRI-5 1988 i.e., differences in mass spawnings (DST= 0.0069;

SWRI-I I985 H, = 0.0370; H, = 0.0440). Furthermore, signifi- SWRI-2 1986 cant heterogeneity in allelic frequencies existed I I I I I among the hatchery groups at six loci: Aut-1 (G = 0986 0988 0990 0992 0994 0996 0998 1000 20.545; 5 d.f.); Aut-3 (G = 43.801; 5 d.f.); Adh (G Genetic Identity = 41.125; 5 d.f); Idh-1 (G = 15.934; 5 d.f.); Pgm-1

Figure 1. Cluster analysis (UPGMA) of genetic identity values among (G = 40.031; 10 d.f.); and Gpi -1 (G = 19 samples of white seabass 50.031; 5 d.f).

102 BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

Genetic identities between hatchery samples were We could detect no consistent geographic, clinal, slightly less than those between wild samples (data or temporal component to the observed genetic var- not shown). However, all but one of the hatchery iation in wild populations of white seabass from the samples clustered together in figure 1. The genetic Southern California Bight region. This result was similarity between the cluster of wild samples and a not unexpected, given the past history of genetic single composite hatchery sample (produced by av- studies on pelagic marine fishes (Gyllensten 1985; eraging allelic frequencies from all hatchery sam- Waples 1987; Hedgecock et al., 1989). In highly ples) was 0.994. mobile species such as white seabass (Vojkovich and Reed 1983), gene flow among localities is apparently sufficient to homogenize the genetic structure. DISCUSSION The significant F,, values and the occurrence of The electrophoretic analysis of wild samples of variant alleles in specific samples suggested that the white seabass revealed more genetic variability than population of white seabass in the study area may was previously reported for this species. Soul6 and not be panmictic, but rather a dynamic mosaic of Senner (unpublished report to California Depart- very similar subpopulations. Hedgecock et al. (1989 ment of Fish and Game) detected polymorphisms and unpublished data) observed an extremely com- only in alcohol dehydrogenase and phosphogluco- plex population genetic structure in northern an- mutase enzyme systems. Levels of heterozygosity chovy; allelic heterogeneity existed between sexes, reported here were slighly higher than the range of locations, and age classes. Campton and Utter values (0.009-0.043) reported for other members of (1987) stated that gene frequencies in subpopulations the Scianidae (Ramsey and Wakeman 1987). Waples may randomly fluctuate around a global mean for (1987) reported a range of heterozygosity values of the population. The random fluctuations may be sta- 0.009 to 0.087 (mean = 0.031) for ten species of tistically significant even with substantial levels of Southern California Bight marine shorefishes that gene flow (Allendorf and Phelps 1981). Waples (1987) display diverse life-history characters. Average het- observed isozyme loci with significant F,, values erozygosity values of white seabass were similar to (i. e., subdivision) in two marine shorefishes with two of the species studied by Waples: the wooly limited dispersal capabilities - the viviparous black sculpin, Clinocottus analis (H = 0.046), and the perch, Embioticajacksoni (average F,, = 0.444), and ocean whitefish, Caulolatilus pinceps (H = 0.049). the wooly sculpin (average Fs, = 0.042). Marine Subpopulations of white seabass appear to be ge- species that Waples judged to have high dispersal netically similar in the Southern California Bight capabilities showed a lower range of F,, values (0- region. However, significant subpopulation differ- 0.028) than we observed in white seabass. entiation was discovered among the samples in 1988, The results of this study have favorable implica- and significant allelic frequency heterogeneity ex- tions for hatchery enhancement programs. Levels of isted between sampling years at Point Loma. We do genetic variability detected here are similar to levels not infer that discrete subpopulations of white sea- in other species to which genetic marking tech- bass exist in the Southern California Bight area. The niques have been applied (Murphy et al. 1983; Seeb reasons for these differences are unclear at present, et al. 1986). Furthermore, we detected several alleles but may be an effect of the rare alleles, nonrandom that could be used as genetic markers to differentiate sampling of the populations, temporal instability of hatchery stocks from wild stocks. Frequencies of allele frequencies, or selection, as well as discrete alleles such as Ldh-3(40 or 1-50), Mpi-(j10), or Aat- subpopulation structure. Hedgecock and Bartley l(115 or 50) could be increased in hatchery popula- (1988) found significant allelic frequency differences tions through selective breeding, and used to quan- between juvenile California halibut from Mission tify hatchery contribution to a target population Bay, San Diego, and adults from Marina del Rey, (Pella and Milner, 1987). and discussed several testable hypotheses to account Hydrologic and zoogeographic data suggest that for genetic differences within a theoretically pan- the Southern California Bight represents a zoogeo- mictic population. Unfortunately, we do not have graphic province bounded by Magdalena Bay, Baja size or age data on the present collections of white California, to the south, and by Point Conception seabass; therefore a more detailed analysis of the to the north (Briggs 1974). The province is charac- causes of genetic heterogeneity, as was done in stud- terized by fish fauna that show little regional genetic ies of northern anchovy (Hedgecock et al. 1989) is differentiation (Waples 1987). Although our initial not possible at present. description of the population genetic structure of

103 BARTLEY AND KENT: GENETIC STRUCTURE OF WHITE SEABASS CalCOFl Rep., Vol. 31,1990

white seabass suggests that brood stock could be less than N because of different reproductive output collected and progeny released at convenient loca- and unequal sex ratio of adults (Allendorf and Ry- tions within the study area, one should proceed care- man 1987). We should point out that the progeny fully before implementing such a hatchery program. groups we analyzed represent a small proportion of Life-history data on spawning patterns, migration, the progeny produced at SWRI. age of maturity, and growth rate must be collected The precise number of adults participating in the throughout the Southern California Bight region to spawns could not be determined, because the geno- determine if the allozyme similarity in the area is types of the brood stock are unknown and because reflected by phenotypic similarity. We do not know spawning is difficult to see. If all adults contributed if mixing the populations of white seabass in the equally to each spawn, we would expect to see the Southern California Bight through transfers of same alleles in each hatchery sample. Because we hatchery-produced fish would cause problems of observed different alleles in successive progeny hybridization of locally adapted stocks and result in groups, we believe that only a few adults are in- outbreeding depression (Altukhov and Salmenkova volved. Mass spawns result when one or two fe- 1987). In addition, we know nothing of the contri- males ovulate, and each female is fertilized by one to bution of subpopulations from the offshore islands. four males (personal observation). Therefore, al- Genetic analyses of samples from offshore islands though a single mass spawn may represent a genetic and samples outside the Southern California Bight contribution of only a few spawners, a series of mass may reveal more genetic differences that may be a spawns, each with different brood stock contribut- consideration in white seabass enhancement efforts ing gametes, may represent a genetic contribution in lower Baja California or northern California. of a large number of brood stock. Furthermore, a The progeny groups produced from a single production hatchery may be able to have several brood stock at SWRI were more different from each tanks of brood stock spawning, thereby increasing other than were the wild populations from separate the effective population size of adults and the genetic locations. Furthermore, the hatchery populations variability of the progeny. In a hatchery enhance- were slightly differentiated from natural popula- ment program for white seabass in the Southern tions. The differences in genetic variability among California Bight region, the genetic profile of prog- hatchery samples are most likely due to different eny from mass spawnings should be monitored over adults contributing to successive mass spawns. The extended periods of time to insure that levels of ge- relative contribution of adult brood stock to mass netic variability are maintained. Although white spawnings may also change over time. For example, seabass brood stock are adapted to current hatchery the fish contributing to the 1986 mass spawn pos- conditions, dominance hierarchy or reproductive sessed the Gpi-Z( - 240) allele. Because this allele senescence in certain individuals may reduce the was not seen subsequently, we presume the adult (or number of adults spawning and thus decrease the adults) made little or no further contribution to the genetic diversity of progeny. progeny samples. Sampling error and small sample Technology now exists for producing large num- sizes may also account for some differences in allelic bers ofwhite seabass in a hatchery environment. But frequency or presence of rare alleles in hatchery the feasibility of enhancing a depressed natural pop- samples. ulation of white seabass is still controversial. The It is important to note that progeny from a single allozymic data presented in this paper address ge- mass spawn have a different genetic profile and may netic considerations associated with hatchery en- have less variability than natural populations; when hancement. Recently developed molecular genetic additional progeny samples are analyzed, the ge- techniques can also be applied to fishery analyses. netic variability of the entire hatchery product is DNA-level polymorphisms from mitochondrial increased. However, there is a limit to the genetic DNA, and DNA fingerprints may reveal additional variability in progeny from a limited number of markers for hatchery fish and may provide a means adults. We observed a 15% decrease in total gene to follow the contribution of individual brood stock diversity in hatchery samples compared to wild (Hallerman and Beckman 1988). We recommend the samples (0.0440/0.0516). A founding population of continued collection of allozyme data from offshore N = 15 individuals (such as in a mass spawning islands and areas outside the Southern California tank) should preserve 1-1/2N or 97% of the varia- Bight, as well as the incorporation of DNA analyses tion of the original population. This relation will to better understand the population genetic struc- only hold if N is the effective population size (NJ. ture of natural and hatchery populations of white Ne of the white seabass brood stock may be much seabass.

104 BARTLEY AND KENT: GENETIC STRUCTURE 0.F WHITE SEABASS CalCOFl Rep., Vol. 31,1990

ACKNOWLEDGMENTS California halibut, Paralirhrhys cn/$oforriic.ris. Calif. Fish Game 74:119- 127. This research was supported by the Ocean Re- Hedgecock, D., E. S. Hutchinson, G. Li, E L. Stone, and K. Nelson. sources Enhancement and Hatchery Program 1989. Genetic and morphometric variation in the Pacific sardine, Sar- within the California Department of Fish and din& sagax caerulea: comparisons and contrasts with historical data and with variability the northern anchovy, Engraulis mordax. Fish. Game. We gratefully acknowledge the efforts of Bull. U.S. 87:653-671. R. E Ford, C. Donohoe, l? Dutton, R. Orhun, and Kincaid, H. L. 1983. Inbreeding in fish populations. Aquaculture commercial gillnetters R. Miller and R. Phillips in 33215-227. MacCall, A. D. 1989. Against marine fish hatcheries: ironies of fishery collecting white seabass. We thank D. Campton and politics in the technological era. Calif. Coop. Oceanic Fish. Invest. L. McClenaghan for computer programs and dis- Rep. 30:46-48. cussions, D. Trusedale for laboratory assistance, Markert, C. L., and I. Faulhaber. 1965. Lactate dehydrogenase isozyme patterns of fish. J. Exp. Zool. 159:319-332. A. Ibarra for the Spanish translation of the abstract, Murphy, B. R., L. A. Nielsen, aiid B. J. Turner. 1983. Use of genetic S. Fox for preparing the tables, and D. Hedgecock tags to evaluate stocking success for reservoir walleyes. Trans. Am. and an anonymous referee for helpful reviews. Fish. SOC.212457-463. Nei, M. 1973. Analysis of gene diversity in subdivided populations. Proc. Nat. Acad. Sci. USA 70:3321-3323. L ITE RATU R E C ITE D . 1977. F-statistics and analysis of gene diversity in subdivided Aebersold, P. B., G. A. Wiiians, D. J. Teel, G. B. Milner, and populations. Ann. Hum. Genet. 441:225-233. F. M. Utter. 1987. Manual for starch gel electrophoresis: a method for -. 1978. Estimation of average heterozygosity and genetic dis- the detection of genetic variation. Springfield, Va.: NOAA Technical tance from a small number of individuals. Genetics 89:583-590. Report NMFS 61, 19 pp. Netboy, A. 1974. The salmon: their fight for survival. Boston: Hough- Allendorf, E W., and S. R. Phelps. 1981. Use of allelic frequencies to ton Mifflin. 613 pp. describe population structures. Can. J. Fish and Aquat. Sci. 38:1507- Pella, J. J., and G. B. Milner. 1987. Use of genetic niarks in stock 1514. composition analysis. In Population genetics and fishery maiiage- Allendorf, E, and N. Ryman. 1987. Genetic management of hatchery ment, N. Ryman and E Utter, eds. Seattle: Univ. Wash. Press, stocks. Irr Population genetics and fishery management, N. Ryman pp. 247-276. and F. Utter, eds. Seattle: Univ. Wash. Press, pp. 141-160. Ramsey, I? R., and J. M. Wakeman. 1987. Population structure of Allendorf, F. W., and E M. Utter. 1979. Population genetics. In Fish Sciaeilop orellartis and Cyrrosriori rrchu/osrrs (Pisces). Copeia 1987:682- physiology, vol. 8, W. J. Hoar, D. J. Randall, and J. R. Brett, eds. 695. pp. 407-454. Ridgeway, G. J., S. W. Sherburiie, and R. D. Lewis. 1970. Polyniorph- Altukhov, Y. P., and E. A. Salmenkova. 1987. Stock transfer relative to isms in the esterase ofAtlantic herring. Trans. Am. Fish. Soc. 99147- natural organization, management, and conservation of fish popula- 151. tions. Iri Population genetics and fishery management, N. Rymaii Rutledge, W. I? 1989. The Texas marine hatchery prograni-it works! and F. Utter eds. Seattle: Univ. Wash. Press, pp. 333-343. Calif. Coop. Oceanic Fish. Invest. Rep. 30:49-52. Briggs, J. C. 1974. Marine zoogeography. N.Y.: McGraw-Hill, 475 pp. Ryman, N., and F. Utter. 1987. Population genetics and fishery man- Campton, D. E., and F. M. Utter. 1987. Genetic structure of anadroni- agement. Seattle: Univ. Wash. Press. 420 pp. ous cutthroat trout (Salmo rlarki rlavki) populations in the Puget Seeb, J. E., L. W. Seeb, and E M. Utter. 1986. Use of genetic marks to Sound area: evidence of restricted gene flow. Can. J. Fish. Aquat. Sci. assess stock dynamics and management programs for chum salmon. 44573-582. Trans. Am. Fish. SOC.115:448-454. Chakraborty, R., and 0.Leimar. 1987. Genetic variation within a sub- Shaw, C. R., and R. Prasad. 1970. Starch gel electrophoresis of en- divided population. hi Population genetics and fishery management, zymes - a compilation ofrecipes. Biochem. Genet. 4:297-320. N. Rymaii and E Utter, eds. Seattle: Univ. Wash. Press, pp. 89-120. Skogsberg, T. 1939. The fishes of the family Scianidae (croakers) of Clayton, J. W,, and D. N. Tretiak. 1972. Amine-citrate buffers for pH California. Calif. Fish Game Bull. 54:l-62. control in starch gel electrophoresis. J. Fish. Res. Board Can. 29:1169- Slatkin, M., and N. H. Barton. 1989. A comparison of three indirect 1172. methods for estimating average levels of gene flow. Evolution Gyllensten, U. 1985. The genetic structure of fish: differences in the 43:1349-1368. intraspecific distribution of biochemical genetic variation between Sneath, I? H. A,, and R. R. Sokal. 1973. Numerical . Sail marine, anadromous and freshwater species. J. Fish. Biol. 26:691- Francisco: W. H. Freeman, 573 pp. 699. Sokd, R. R., and E J. Rohlf. 1981. Biometry. New York: W. E Freeman, Hallerman, E. M., and J. S. Beckman. 1988. DNA-level polymorph- 859 pp. isms as a tool in fisheries science. Can. J. Fish. Aquat. Scl. 45:1075- Vojkovich, M., and R. J. Reed. 1983. White seabass, Arvarrosriori riohi/is, 1087. in California-Mexican waters: status of the fishery. Calif. Coop. Harris, H., and D. A. Hopkinson. 1976. Handbook of enzyme electro- Oceanic Fish. Invest. Rep. 24:79-83. phoresis in human genetics. Amsterdam: North Holland Waples, R. S. 1987. Multispecies approach to the analysis of gene flow Publishing Co. 281 pp. in marine shore fishes. Evolution 41 :385-400. Hedgecock, D., and D. M. Bartley. 1988. Allozyme variation in the Wright, S. 1943. Isolation by distance. Genetics 28:114-138.

105 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

THE VERTICAL DISTRIBUTION AND FEEDING HABITS OF TWO COMMON MIDWATER FISHES (LEUROGLOSSUSSTILBIUS AND LEUCOPSARUS) OFF SANTA BARBARA

GREGOR M. CAILLIET ALFRED W. EBELING Moss Landing Marine Laboratories Department of Biological Sciences P. 0. Box 450 and Marine Science Institute Moss Landing, California 95039 University of California Santa Barbara, California 93106

ABSTRACT adaptaciones morfol6gicas que le permiten alimen- Leuvoglossus stilbius (Bathylagidae) was abundant tarse por succidn y predar sobre organismos pe- in the nearshore Santa Barbara Basin (SBB), but less quefios y poco activos. A diferencia, la especie S. so in the more offshore Santa Cruz Basin (SCB). leucopsarus est6 mejor adaptada a una alimentaci6n Stenohvachius leucopsarus (Myctophidae) was abun- activa, predando sobre presas mPs grandes y mis dant in both basins. L. stilbius is adapted morpho- escapadizas. L. stilbius se aliment6 principalmente logically to feed by suction and to eat smaller, less de noche en aguas superficiales de la SBB. Durante active organisms. S. leucopsarus is better adapted to todo el aiio el alimento consisti6 de apendicularias y feed by grasping, and to eat larger, faster, and more salpas, reflejando la persistente disponibilidad de es- elusive prey. In the SBB, L. stilbius fed, mostly at tos organismos a lo largo del ciclo anual. El regimen night in surface waters, on larvaceans and all alimenticio fue similar en la SCB, si bien la dieta year, reflecting the seasonally consistent abundance vari6 de acuerdo con la variaci6n estacional en la of these prey items. In the SCB, it fed less intensely, abundancia de las presas. No se observaron diferen- mostly at night in surface waters, and its diet varied cias cronol6gicas marcadas en 10s hibitos alimenti- with the seasonal abundance of its gelatinous prey. cios de S. leucopsavus. La dieta estuvo compuesta S. leucopsarus fed mainly on crustaceans, and it did principalmente por crusticeos, en especial de eu- not exhibit a distinct feeding chronology. It ate sim- faQsidos cuando istos eran mis abundantes. El ilar prey all year in both basins, but euphausiids alimento fue similar en ambas cuencas. Por consi- dominated the diet when they were most abundant. guiente, L. stilbius es una especie adaptada a1 am- Thus L. stilbius is well adapted to inshore, eutrophic biente costero eutr6fico donde puede alimentarse de midwater habitats, where it can easily eat abundant apendicularias y salpas en abundancia, y que debido larvaceans and salps, and may act as a trophic link a sus migraciones verticales difusivas es probable between the shallow gelatinous zooplankton and the que represente el eslabon trdfico entre el zooplanc- deep sea through its diffuse vertical migrations. ton gelatinosos de aguas poco profundas y las aguas Offshore, its primary food is less dense, only sea- oceinicas profundas. Lejos de la costa, su alimento sonally available, and restricted to surface waters. principal est5 menos densamente distribuido, y so- S. leucopsuvus is better adapted to eat the more var- lamente disponible estacionalmente y restringido a ied food resources offshore. Its cohesive pattern of las aguas superficiales. La especie L. leucopsartis esti vertical migrations takes it into the food-rich sur- mejor adaptada a un regimen alimenticio oceinico face waters at night, where it consumes crustacean mPs variado. Su patr6n de migraci6n vertical cohe- prey and trophically transports their calories to the sivo le permite predar en aguas superficiales ricas en deep sea. alimento (donde consume crusticeos) y transportar las calorias hacia las aguas profundas. RESUMEN La abundancia de Letivoglosstis stilbius (Bathylagi- INTRODUCTION dae) fue mPs alta en la cuenca de Santa Barbara (SBB) According to Lavenberg and Ebeling (1967) “the que en la cuenca de Santa Cruz (SCB), ubicada a diversity of the mesopelagic and bathypelagic faunas mayor distancia desde la costa. La abundancia de increases with vertical expansion of their habitats Stenobrachiiis leucopsanis (Myctophidae) fue similar offshore. ” This trend is evident when one compares en ambas cuencas. La especie L. stilbius presenta the fish faunas of two deep-sea basins off Santa Bar- bara, California (Ebeling et al. 1970a; Brown 1974). The Santa Barbara Basin (SBB), located inshore of ~~ ~~~ [Manuscript rece~vedFebruary 2,1990 ] the Channel Islands, is relatively shallow (600 m,

106 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

with a 425-tn sill), generally isolated froin other ba- lates the sizes of the inshore and offshore populations sins, and enriched by coastal runofc it has a rela- of these species? Of the four key factors listed by tively simple but abundant epipelagic and upper MacArthur and Connell (1966) that regulate popu- mesopelagic fish fauna. In contrast, the offshore lation sizes (reproduction, migration, mortality, aiid Santa Cruz Basin (SCB) is much deeper (>2000 m) food resources), we decided to evaluate the fourth. and is outside the immediate coastal influence, in We investigated the morphological adaptations, closer contact with the deeper oceanic environment; feeding habits, and vertical migration patterns of the it contains a more diverse but less abundant fish two fishes to determine what kinds of prey they fauna. Here, allochthonous species increase in rela- took, how they migrated vertically relative to prey tive abundance with large-scale seasonal changes in availability, how their use of available food might water-mass types, and bathypelagic species occur influence their relative success in the inshore and off- below 500 m. shore areas, and how they might affect the vertical Off California, evidence suggests that phyto- flux of organic material in the water column. plankton production and standing crop are highest inshore (Malone 1971), and the zooplankton diver- sity increases as zooplankton density decreases MATERIALS AND METHODS offshore (Longhurst 1967). The inshore SBB is re- Fishes were collected from 1964 through 1968 garded as highly productive (Emery 1960; Soutar during 33 regular cruises of the RIV Swan off Santa and Isaacs 1969; Sholkovitz and Gieskes 1971). Ebe- Barbara, California (cf Ebeling et ai. 1970a; Brown ling et al. (1970a) reported that catch volumes of 1974). All collections were made with a 1.8-in fishes were higher there, whereas volumes of inver- IKMT, which had a lining of 1-cm stretch mesh tebrate micronekton did not differ significantly be- netting, followed by a standard zooplankton net and tween basins. The variability in the offshore catch a cod-end sampler, divided into four chambers by was greatly influenced by seasonal invasions of electronically closed gates (Aron et al. 1964; Bour- bulky organisms like salps. If salps are eliminated beau et al. 1966). Samples of animals from particular from the analysis, invertebrate standing crop was depth intervals were thus separated by the sequen- also greatest in the SBB. tially closed gates. The trawl’s spreader bar con- The California smoothtongue, Leuvoglossus stil- tained electronic sensors to monitor depth aiid water bius, (Bathylagidae) and northern lampfish, Steno- temperature. A flowmeter measured sampling ef- brachills leucopsartis, (Myctophidae) are the dominant fort in meters trawled. The signals from all sensors fishes in the midwater community of animals that is and flowmeter were transmitted simultaneously especially abundant off southern California (Ahl- through the towing cable to shipboard recorders. strom 1969; Ebeling et al. 1970a, 1970b; Brown Collections were regularly made in the generally 1974). L. stilbius was ranked first, and made LIP 58% recognized major depth zones: (1) epipelagic, in the of all fishes sampled. Its catch rate averaged 6.1 surface wind-mixed layer about 0-200 in; (2) upper adults per kilometer flow through an 1.8-in Isaacs- mesopelagic, within the permanent thermocline, Kidd midwater trawl (IKMT) in the SBB. It ranked about 200-400 in; and (3) lower mesopelagic, in the only third, 16%, and 0.4 km-’ in the SCB. Like- dysphotic depths, below 400 in. Maximum trawl wise, S. leucopsalus ranked second, and had abun- depth varied between 500 aiid 1000 ni, depending dances and catch rates of33% and 5.4 km-’ inshore, on the area sampled. Only shallow and mid-depth and ranked fifth, and had values of 11% and 0.9 samples could be taken in SBB, which lacks a bathy- km-’ offshore. Farther offshore, the numbers of pelagic zone. The more typically oceanic and deeper both species dwindle, but S. leucopsanis is a bit more SCB has a bathypelagic zone that extends well be- abundant than L. rtilbiiis (Pearcy 1976; Willis and low the depth of the lower mesopelagic. Samples Pearcy 1980). Although the distributional centers of were taken throughout the year at night and during both fishes occur off California, L. stilbius ranges the day. A total of 205 stations yielded 631 collec- from Alaska to the Gulf of California, while S. leu- tions, 363 of which were from discrete-depth hauls copsavtis occurs all the way from the Bering Sea to whose vertical excursions were at least 50% within the tip ofBaja California (Miller and Lea 1972; Hart the 200-in depth intervals. An additional 40 samples 1973; Eschmeyer et al. 3 983). through broader depth intervals were used to esti- Ebeling et al. (1970b) theorized that “among me- mate seasonal abundance of potential prey, such as sopelagic fishes L. stilbitis and to a lesser extent salps and euphausiids. S. leucopsants may best exploit the rich inshore basins All fishes were preserved on board in 10% buff- of the borderland. ” What, then, differentially regu- ered Formalin and seawater, and subsequently

107 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

changed to 45% isopropyl alcohol ashore. Speci- significantly alter feeding habit results. However, mens of L. stilbius and S. leucopsavtis were routinely Lancraft and Robison (I 980) found simulated prey identified, counted, and measured, along with other items in 23.3% of the guts of S. leucopsavus. There- fishes and invertebrates of each trawl catch within a fore, we used stomach contents with scales to esti- few months after collection (Ebeling et al. 1970a). mate possible bias in this species. We compared rank Small fishes (less than 50 mm standard length, SL) orders of diets of fish with scales in their stomachs were arbitrarily distinguished from larger fishes. with those without scales for both basins, using Very small individuals of both species were caught Kendall’s nonparameteric rank correlation (Sokal but were not adequately sampled because many and Rohlf 1969). probably escaped through the mesh. Abundances of With contents intact, the fullness of the gut was both species were standardized among collections subjectively scored as: 0 = empty; 1 = 25% full; 2 by sampling effort, measured in meters but defined = 50% full; 3 = 75% full; and 4 = 100% full. The in units of kilometers towed. contents were then removed, identified to the lowest The depth distributions ofthe two species, pooled possible taxon, measured with an ocular microme- for all months, were analyzed separately for four ter, and counted. The percent volume contribution time intervals - late night (0001-0600 hrs); morn- of each prey item was subjectively estimated. Any ing (0601-1200 hrs); afternoon (1201-1800 hrs); and intestinal parasites were identified, counted, and early night (1801 -2400 hrs) - and for five depth in- measured. tervals - 0-200 m, 201-400 m, and >400 m in the The index of relative importance (IRI) of each SBB and SCB, plus the deeper zones 600-800 m and prey item was estimated for food-containing fish by >800 m in the SCB. Histograms of bathymetric each time and depth category as a linear combination distribution of abundances were constructed for of its numerical importance (N),volumetric impor- these four categories of day and night captures in the tance ( V),and frequency ofoccurrence (FO) (Pinkas “at-depth” hauls. The same procedure was followed et al. 1971). The numerical importance of a particu- for analyzing the depth distributions of two species lar item was the percentage ratio of its abundance to of prey caught in the trawls. Their seasonal abun- the total abundance of all items in the contents. Its dances, pooled for all years by month and season, volumetric importance was its average percent vol- were also analyzed for both basins. ume. Its percent frequency of occurrence was the Measurements and counts of the following ali- percentage of fish containing at least one individual. mentary structures were taken to compare the feed- The combination resulted in: IRI = (N + V)x F, ing abilities of the two species: jaw length; width of which is represented by the area of a rectangle re- gape; number and size of teeth; number, size, and solved by plotting the three importance measures on structure of gill rakers; and the length and general a three-way graph (Cailliet et al. 1986). The value of structure of the stomach, intestine, and pyloric IRI ranges from zero, when all three values are zero, caeca. In all, 647 L. stilbiiis and 677 S. leticopsavtis to 20,000, when all three indices are 100% (a mono- were measured, weighed, and dissected. The diet). The IRI ranks the relative importance of di- lengths and weights were used to calculate a “con- etary items, and the three-way graph indicates dition factor” (Cailliet et al. 1986). The gut was ex- which measures of importance were most mean- posed by opening the coelom. The entire alimentary ingful. tract was removed by cutting at the esophagus and Stomach contents of the two fishes were com- pulling it out. The stomach was then split longitu- pared between basins and among oceanographic pe- dinally from the pyloric sphincter to the esophagus. riods within basins. These periods were defined as: For L. stilbius, which has two stomachs, the cut (I) a surface mixing period from January through was made from the posterior end of the pyloric April, when cold weather and stormy turbulence stomach, through the cardiac stomach to the esoph- causes surface cooling; (11) an upwelling period of agus. Only the contents of the pyloric stomach were surface enrichment from May through July, when analyzed for this study, since the cardiac stomach the California Current is strongest and intensifies contents were more likely to reflect net feeding (An- the counterclockwise current gyre over the southern derson 1967; Lancraft and Robison 1980). California continental borderland; and (111) a period For S. leucopsanis, however, there was only one of thermal stratification, from August through De- stomach to analyze. Collard (1970) felt that net feed- cember (Brown 1974; Jones 1971; Sholkovitz and ing was minimal in his study of s. leircopsavtrs feed- Gieskes 1971). ing habits. Likewise, Hopkins and Baird (1975) Possible correlations of rank hierarchies of stom- reported that feeding in midwater trawls did not ach contents were tested between species, basins,

108 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

and among oceanographic periods via Kendall's tau rank correlation (Sokal and Rohlf 1969). We used the numerical importance of prey items eaten (lowest possible taxon, but not necessarily species) to calcu- late several diversity indices. Because all indices pro- duced similar results, we will present only the NM (number ofmoves) indices (Fager 1972). The differences in abundance of two of the prin- cipal food items among oceanographic periods were also estimated. We calcualted the mean numbers of Salpafusijormis and Euphausiapacijica per km trawled and compared them with the seasonal occurrence of these prey in the stomachs of L. stilbius and S. leucop- sarus. This analysis could not be done for smaller or delicate items, such as larvaceans, , and , because of destruction, avoidance, es- cape, or extrusion. To determine recency and amount of feeding for different time and depth categories, we combined estimates of fullness and state of digestion. A 4 X 4 matrix of fullness by digestion for each time and depth category resolved major feeding states of (A) not recently eaten or full, including empty stomachs (i.e., fullness states 0, 1, and 2 vs. digestion states 1 and 2); (B) recent but not full (fullness states 0, 1, and 2 vs. digestion states 3 and 4); (C) recent and full (fullness states 3 and 4 vs. digestion states 3 and 4); and (D) full but not recent (fullness states 3 and 4 vs. Figure 1. Mouth shape and front view of the head of L. stilbius (above) and S. leucopsarus (below). digestion states 1 and 2). Fullness-recency histo- grams measured frequencies of the major feeding states for both species between basins among the four time intervals (late night, morning, afternoon, and early night) and the three depth intervals (0-200,201-400, and >400 m).

RESULTS

Alimentary Morphology L. stilbius has a smaller mouth than S. leucopsarur (figure 1). The mean ratio of upper jaw length to standard length was 6.2 (n = 170) for adult L. stil- bitis, but 17.3 (tz = 41) for adult S. leucopsavtrs. The mean ratio ofgape width to standard length was 5.9 for L. Stilbiars and 8.9 for S. letlcopsarus (figure 2). Assuming that these fishes can open their mouths to a 45" angle, the effective mouth area of S. leucopsartis is about four times that of L. stilbius. L. stilbius has very few teeth in its mouth (figure 1). It has none on its premaxilla or tongue and few on its palatine, vomer, and dentary bones (Chapman 1943; Borodulina 1968). S. leucopsavus, on the other -lOmm hand, has well-developed premaxillary, palatine, Figure 2. Side views of heads and mouths of a fish with a small mouth and L. pterygoid, and dentary teeth (Bolin 1939; Jollie stilbius (above), and a fish with a large mouth and S. leucopsarus (below).

109 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES ColCOFl Rep., Vol. 31,1990

1954; Berry 1964). Unlike L. stilbiiu, it has well- The number of pyloric caeca and relative intes- developed pharyngeal teeth to help move food tinal lengths also differ considerably (figure 5). into the gut Uollie 1954). With more teeth, S. leir- L. stilbius has more caeca (8-11, cf. Borodulina copsarus may better grasp and hold larger and more 1968) than S. leucopsartis (4-6, cf. Jollie 1954). Its active prey. intestine length averaged 50.5% of its SL (n = 22), L. stilbius has significantly more gill rakers on its compared with only 28.0% for S. leucopsarus (n = first arch (26-29, cf. Borodulina 1968) than S. leu- 35). copsariis (17-19, cf. Jollie 1954; figure 3). The front edges of its rakers are smooth, not toothed like those of S. leucopsartis. The average distance between rak- ers of an individual measuring 60 mm SL was only 0.3 mm, compared with 0.7 mm for S. leucopsariis. Also, the gill rakers of L. stilbius are more broadly flattened (Borodulina 1968) and tend to close to- gether when water is forced over them. The stomachs of the two fishes also differ mark- edly. L. stilbiiis has a double stomach, with a cardiac portion preceding the pyloric portion (figures 4 and 5). The cardiac stomach is covered with a black pig- ment, an adaptation that may prevent light from bioluminescent prey from showing through (Mc- Allister 1961). This stomach has a very thick wall, and its inner mucosa is made up of many posteriorly 10mr , oriented rugae (figure 5). The pyloric stomach is Figure 4. Lateral cutaway view of the body and alimentary tract of L. stilbius thin-walled and flexible. In contrast, S. leircopsartis (above) and S. leucopsarus (below). has only one thick-walled stomach, also covered with a black pigment (McAllister 1961). The inter- nal mucosa is made up of typhlosole ridges that run longitudinally (figure 5).

STOUIC" ITE 5 I #NE 216XSL 28 ox SL

Figure 5. Internal alimentary structures of L. stilbius (above) and S. leucop- sarus (below). The upper left drawing for each species is the alimentary canal, showing the stomach@) (stippled and starred), intestinal tract, and pyloric caeca. The upper right drawing for each species shows a longitudinal cutaway of the stomach, either lined with posteriorly oriented rugae (L. stil- bius) or typhlosole ridges (S. leucopsarus). Bar diagrams indicate the aver- Figure 3. Lateral cutaway view of the gill arches and rakers of L. stilbius age lengths of stomach(s) and intestine relative to standard length for both (above) and S. leucopsarus (below). species.

110 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

Feeding Habits ceans were both numerically and volumetrically In the SBB, L. stilbius ate primarily larvaceans important, while the salps were important only (genus Oikopleuvu) and salps (probably Thaliu demo- volumetrically. In both basins the prey diversity in- cvatica and s.fusqovmis: Berner 1957, 1967; Hubbard dices (Fager 1972) were low (SBB, NM = 0.20 and and Pearcy 1971; M. Silver, pers. comm.), followed, SCB, NM = O.ll), indicating that L. stilbitir con- in order, by ostracods, small copepods (1-2 mm), centrated on only a few prey items, hence the trun- zoea, and E. pacifica (figure 6). Salps, which were cated shape of the IRI diagrams (figure 6). Net larger than the more numerous larvaceans, made up feeding could not have biased these results, which the greater dietary bulk. Less-important items in- were based on pyloric stomach contents only. In- cluded copepods (2-3 mm and <1 mm), chaetog- deed, fish scales were never found in these stomachs. naths, fish eggs, siphonophores, nauplii, and larger The diet of S. leucopsarus was also similar between copepods (>3 mm). basins (tau = 0.63; P << 0.01) but was uncorrelated The rank order ofprey in L. stilbius stomachs was with that of L. stilbius (SBB: tau = 0.16, P - 0.35; significantly correlated between basins (Kendall’s SCB: tau = 0.08, P- 0.64). IntheSBB, S. leucopsartis tau = 0.51, P << 0.01). Even so, the SCB fish had ate ostracods, E. pacifica, and a variety of “large” a slightly more varied diet, which differed in minor copepods, with no item predominating unless all ways. They ate relatively fewer ostracods, more size classes of copepods are pooled (figure 7). Less- nauplii, and more large copepods, with such things important items were fish eggs, the euphausiid Ne- as amphipods (Hypevia gulba) and shrimp (mysids matoscelis dqjcilis, zoea, the amphipod H. galba, and sergestids) in lesser amounts. Also, the larva- shrimp (mysids and sergestids), chaetognaths, fish larvae, siphonophores, salps, and “small” copepods (<1 mm). The SCB S. leucopsavus ate relatively more eu- ,~Oik “1 ”’ phausiids (figure 7). Like L. stilbicrs, they ate rela- b 40 tively fewer ostracods, and more large copepods. D n=256 5 Amphipods were mainly Pavaphvonima cvassipes. The s 20 prey diversity inshore (NM = 0.28) was lower than in fish offshore (NM = 0.34), but in both cases was 0 higher than for L. stilbiur, implying that S. leucopsa- ! YMS generally ate more types of food, hence the elon- 2 20 gated appearance of the S. leucopsarus IRI diagrams 8 cop>3 (figure 7). 4OJ Fish scales occurred in relatively high frequencies in S. leucopravus stomachs (26.5% of SBB and 13.1O/O

n=129 6or””l 401 SBB n=349

o- 10 20 30 , , , , , % F.O. . a,.

40 U sa cop1

Figure 6. Percent composition of major prey items in number, volume, and frequency of occurrence (% F.O.) and ranked by the index of relative impor- tance (IRI) from left to right for L. sfilbius in the Santa Barbara Basin (SBB) and Santa Cruz Basin (SCB). n = the number of guts analyzed. Code of abbreviations: Ch = chaetognaths; Cop <1 = copepods smallerthan 1 mm; Cop 1-2 = copepods 1-2 mm; Cop 2-3 = copepods 2-3 mm; Cop >3 = copepods larger than 3 mm; CrD = crustacean debris; Ep = Euphausia pacifica (euphausiid); FE = fish eggs; FL = fish larvae; FS = fish scales; Hg = Hyperia galba (amphipod); na = nauplii; Nd = Nematoscelis difficilis (euphausiid); Oik = Oikopleura spp. (larvacean); Ost = ostracods; Pcr = Paraphronima crassipes [amphipod); Rad = radiolarian; Sa = salps; Sh = shrimp (mysids and sergestids); Si = siphonophores; 20 = zoea; U = Figure 7. Percent composition of major prey items for S. leucopsarus. Abbre- unidentifiable. viations as for figure 6.

111 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

of SCB fish). These scales, an unlikely food, were sonal changes in the food supply. The fish ate mostly probably ingested in the trawl net as “net feeding” larvaceans and salps during periods I and 11, but (Collard 1970; Hopkins and Baird 1975; Lancraft mostly copepods during period 111 (figure 8). Ranks and Robison 1980). However, “scaled” and “un- of dietary items were not significantly correlated scaled” diets were significantly correlated (SBB: tau betweenperiodsIandI1 (tau = 0.18, P-O.4), mainly = 0.83, P << 0.001; SCB: tau = 0.92, P << because larvaceans did not dominate in period I1 and 0.001). Therefore, net feeding apparently did not salps did not even rank in the top six during period systematically bias the observed dietary composi- I. Periods I1 and 111 were also not correlated (tau = tion, and fish with scales in their stomachs were not 0.45, P - 0.02) because copepods of all sizes were eliminated from the analysis. commonly eaten during period 111, but larvaceans and salps were not. Of secondary rank during period I were large and small copepods, crab zoea, and Seasonal Variation in Feeding Habits chaetognaths. During period 11, salps, small cope- In the SBB, the prey of L. stilbius reflected the pods, euphausiids, and zoea were secondary. During relatively even yearly distribution of the food sup- period 111, a diversity of large and small copepods ply. Fish ate about the same kinds prey all year, of outranked zoea, larvaceans, and salps. SCB catches mostly larvaceans and salps (figure 8), and the ranks of S. fusijovmis (unlike those from the SBB) de- of food items were significantly correlated among creased abruptly from period I to periods I1 and 111 seasons (W = 0.85, P << 0.001). Salps were equally (figure 9). abundant in the cold mixing (I) and upwelling (111) In the SBB, the relatively varied diet of S. leucop- periods (figure 9), but were lower during the spring suvus did not reflect the seasonal changes in food upwelling season (11), which may account for a supply. S. leucopsartis ate mostly small copepods and slight coincident change in the fish’s diet. Although larvaceans ranked first in dietary importance during periods I and 11, salps ranked first during the warmer stratification season (111), when they were slightly Salpa fusiformis more common in the inshore plankton. During pe- riod 111, ostracods ranked second, and larvaceans third in the fish’s diet. We have no way of assessing 40i the availability of larvaceans like Oikopleuva. Eu- phausiids were eaten in noticeable numbers only 5c ””1nn during period I, followed in rank by large copepods, LU crab zoea, and small copepods. During periods I1 10 and 111, ostracods, small copepods, and large cope- pods completed the diet. n In the SCB, however, significant changes in the n=52 n=67 n=95 diet of L. stilbius seemed to reflect concomitant sea- 60

n=92

40 tc e 30 aP 40 C SCB n=56 n=39 n=34 20

10

n u Jan- Apr May-Jul Aug-Dec I II 111 Jan-April May-Jul Aug-Dec I II 111 Figure 9. Changes in abundance (number per km sampled) of Salpa fusifor- mis, a major prey of L. stilbius, from IKMT discrete depth tows (n = sample Figure 8. Percent composition of the six top-ranking prey items for L. stilbhs size) in the upper 500 m of the Santa Barbara (above) and Santa Cruz for three oceanographic periods. Abbreviations as for figure 6. (below) basins for three oceanographic periods.

112 CAILLIET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

ostracods during period I, but more euphausiids Euphausia pacifica (E. paci_fca) during periods I1 and I11 (figure 10). Ranks of food items differed significantly between periods I and I1 (tau = 0.37, P - 0.045), but were correlated between periods I1 and 111 (tau = 0.45, P < 0.01). E. pacijica catches were low during period I (figure ll), when the fish ate mostly copepods and ostracods. E. paci$ca ranked first in the diet during periods I1 and 111, but was much more abundant in catches during period I1 than 111. The remainder of the fish's diet comprised large copepods, euphau- siids, and chaetognaths during period I; large and small copepods, ostracods, and fish eggs during pe- riod 11; and ostracods, small copepods, other eu- phausiids (N. dfficilis), and large copepods during period III. ''01 scB In the SCB, however, the dominant prey of S. leucopsarus were also the most abundant in midwater sE '"1 trawl catches. S. leucopsarus ate both large and small K copepods during period I, when E. pacijica was not 100 abundant, and E. pacijcu during periods I1 and 111 (figure lo), when they were abundant in the plank- ton (figure 11). Ranks of food items differed mark- n U edly between periods I and I1 (tau = 0.20, P - 0.3), Jan- Apr May-Jul Aug-Dec and between periods I1 and 111 (tau = 0.49, P - 0.04), I II 111 even through E. pac$ca dominated the diet in these two periods. The remainder of the diet comprised Figure 11. Changes in abundance of Euphausia pacifjca, a major prey of S. leucopsarus, from IKMT discrete depth tows (n = sample size) in the upper the same kinds of items eaten in the SBB. 500 m of the Santa Barbara (above) and Santa Cruz (below) basinsfor three periods. Diel Vertical Migrations L. stilbius migrated vertically in a diel pattern that differed somewhat between basins. In the SBB, the fish were abundant in the surface waters during the afternoon and early night, in deep waters during late Santa Barbara Basin night, and at mid-depth during the morning (figure Leuroglossus stilbius Time of Day (hours) 12). A significant portion of the population was 0001-0600 N211 ~ , ~ f-2400 0 200 ;, i - E z 201-400 141 111 -E -v) 401-600 e)-( 1201 1151 n=83 e -E 0 (0 20 10 a0 " 10 20 30 40 0 10 20 10 40 0 10 20 ,o 40 5 Sfenobrachius leucopsarus P 1 5 201.400 111

401-600 1201 115) ;0 , , ,(; , ,,:, o.200k0 10 20 30 40 I',;0 10 20 30 40 0 10 20 30 40 0 10 20 10 40 Mean Number per km (?%E.) ,:; p

Figure 12. Diel vertical distribution patterns for L. stilbius and S. leucopsarus Jan-April May-Jut Aug-Dec in the Santa Barbara Basin. Data were pooled for all months among four 6- I II 111 hour time intervals and three 200-m depth intervals. The horizontal axis mea- sures abundances, t standard errors, standardized by trawling effort as Figure 10. Percent composition of the SIX top-ranking prey items for S. leucop- numbers per km flow. The numbers in parentheses represent the sample size saius for three oceanographic periods. Abbreviations as for figure 6. (number of trawls) for each time-depth category.

113 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

00M-0600 0601-1200 l2M-1800 180,-2400 found near the bottom of this shallow basin during 0-200a all periods, and the fish did not stay near the surface 100 all night. In the deeper SCB, they had a broader I vertical distribution: some of the fish were found in the surface waters in the evening, especially during 20 ...... the late night and early morning, and more were 1401 153) E O 201-400m found in deeper water (401-600 m), especially dur- ing late night and daytime (figure 13). A significant portion of the population occurred at mid-depths - n (<400 m) during all periods...... The migratory pattern of S. leucopsurus, on the ...... (uI1 other hand, was quite predictable and similar be- g wont ‘0°1 tween basins. In both the SBB and SCB, most of the population was found in the surface waters at night and at mid-depth during the day (figures 12 and 13). 40::n n n n ...... Consequently, the shallow SBB did not seem to 121) I301 compress the vertical range of S. leticopsnvus like it ABCD ABCD ABCD ABCD did that of L. stilbius, nor did a significant portion of Figure 14. Fullness and recency-of-feeding histograms for L. sfilbius in the the S. leucopsurus population occur below 400 m in Santa Barbara Basin pooled over all months among four 6-hour time and three depth intervals. The vertical axis measures the percent frequency of either basin. occurrence by time-depth category for fullness-recency states: A (open), not recently eaten or full; B (stippled), recent but not full; C (shaded), recent and full; and D (hatched), full but not recent. The numbers in parentheses equal Fullness and Recency of Feeding Relative to Vertical the number of fish in each time-depth category. Migration In the SBB, L. stilbius apparently fed most in- some during both day and night at mid-depth (fig- tensely during the night in the surface waters; it fed ure 14). Fish caught at night in the surface waters and at mid-depth had the highest percentages of “re- cently full” stomachs. Fish caught in the surface dur- Santa Cruz Basin ing the daytime, and below 400 m at all times had Leuroglossus stilbius Time of Day (hours) Owl-0600 0601.1200 1201-1800 1801-24W very high percentages of “not recent or full” stom- - achs, and therefore had not been actively feeding. 0~200 1391 Fish from the mid-depth during the day tended to have contents equally distributed over the fullness- 201-400 (171 , digestion categories, indicating that their stomachs 401-600 1141 1, ;, contained previously ingested items mixed with E EN newly ingested ones. 601-800 151 1171 E In the SCB, however, L. stilbiiis generally fed less - intensely, and mostly at night. Fish had relatively + , I ,!41 , , , ,131 , , , , 101 , 1 1 I high percentages of “recent but not full” stomachs 15 - 0 5 10 15 20 0 5 10 15 20 0 5 10 15 20 0 5 10 15 20 only during the night near the surface, and of “full 5 Sfenobrachius leucopsarus and recent” stomachs only during late night and at

0-200 I391 all depths (figure 15). Few fish fed to fullness, and fish had high proportions of “not full or recent” 201-400 stomachs in all time-depth categories for which there were sufficient samples. 401-600 (141 S. jiisfornzis, a common prey of L. stilbiw, was

601-800 (51 mainly limited to the upper 200 m, whenever it was collected (figure 16). In the SBB, this was rela- 801 + (4) I 161 tively abundant in these surface waters at all times . ,- of the day. Only during the late night was it available -0 5 10 15 20 5 io 15 20 0 5 10 15 20 0 5 10 15 20 Mean Number per km (S.E.) in deeper (401-600 m) waters. In the SCB, however, these salps were caught mainly at night in the upper Figure 13. Diel vertical distribution patterns of the two species in the Santa Cruz Basin. Note the difference in the horizontal axis dimensions. All other 200 m, and they were seldom caught during day- details as in figure 12. light hours.

114 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES ColCOFl Rep., Vol. 31,1990

0001-0600 0601-1200 IZM-1800 1601-2400 0001-0600 0601-1200 1201-1600 1801-2.00 0-zoom 0-zoom 100 1 loo80 n

20

1.01 (31 101 : 201-4001 p 100

n k 40 n f 201 1 y ...... DO (161 ...... 1.01 (15) P 4oom+ 100, n n II n 60

20 L (51) r,...Liw1I L 150) A8CO 46 4i 4BCO 4BCO ABCD ABCD

Figure 15. Fullness and recency of feeding histograms for L. stiibius in the Figure 17. Fullness and recency of feeding histograms for S. leucopsarus in Santa Cruz Basin. All details as in figure 14. the Santa Barbara Basin. All details as in figure 14.

0001-0600 06M-1200 IPOI-I~~ Isa- moo SBB 0-zoom Salpa fusiformis Time of Day (hours) 0001-0600 0601-1200 1201-1800 1801-2400 -E0 , , (N, p, 201-400o.zoor, F=p (!, E 401-600 (20) (15) -. m-4001 -u) - :: 100 m 0 20 40 60 80 100120140 0 20 40 50 80 100120140 0 20 40 50 80 100 0 M 40 60 2 g Bo[ a, -‘E SCB g 60

a, ;20 :4oLDo ...... I 1311 0 201-400401-6000-2OOT(iI, , , , ;, L,,, 5 Y hl,) - ,I, won. 1 ln ; 6 601-800 ,I, ,I, n 801 + 0 20 40 60 8C 100120140 0 20 40 60 80 100120140 0 20 40 60 80 100 0 20 40 60 ::I 1- .... 1 h.... 0 :::: (,,) ...... (191 I51 Mean Number per km (S.E.) 4sco .BCO ABCD ABCO

Figure 16. Diel vertical distribution of patterns of Salpa fusifofrnis, a major Figure 18. Fullness and recency of feeding histograms for S. leucopsarus in prey of L. stilbius, in the Santa Barbara (above) and Santa Cruz (below) the Santa Cruz Basin. All details as in figure 14. basins. The horizontal axis measures the abundance standardized for trawl- ing effort as numbers per km flow, * standard errors. Data were pooled for all months for four 6-hour time intervals and three or five 200-m depth inter- centages of “not recent or full” stomachs, indicating vals. The numbers in parentheses represent the number of IKMT tows In each that they had not eaten much at depth. time-depth category. In both basins E. pacifica, a common prey of S. leucopsavus, exhibited a typical migration pattern In both basins, S. leucopsanis appeared to have fed of occupying the upper 200 m during the night, and at all times and depths as the opportunity arose. dwelling mainly between 200 and 400 m during the It did not exhibit a distinct feeding cycle. High day (figure 19). This species was abundant in both percentages of “recent but not full” and “full and basins but did not appear in significant numbers in recent” stomachs occurred in most time-depth cat- water deeper than 400 m. Therefore, S. leucopsants egories (figures 17 and 18). However, the highest must have consumed these euphausiids in the upper percentages of recent feedings were observed in fish 400 m, no matter what time of day. from the upper 400 in in both basins. Fish from L. stilbius was more heavily parasitized inshore, waters deeper than 400 m generally had high per- whereas S. leucopsavus appeared equally parasitized

115 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

SBB houses and locate the animal either by the beating of Euphausia pacifica Time of Day (hours) 0001-0600 0601-1200 1201-1800 1801-2400 its tail or from created by organ- isms living on or in contact with its house. They can then suck the larvacean out, ingest the whole com- plex, or scare the animal away from its house and v 115) 2 401-600 then catch it and suck it in. Salps are patchy and seasonally common, and can exist solitarily or in strings (Berner 1967; Hubbard and Pearcy 1971; Silver 1975). They are probably

201-400 encountered by individual fish, presumably in the surface waters and sometimes in the daytime when ln L. stilbius can see them and suck them in. Both ge- 5 601-800 latinous prey were often found in quantity in an 801+ individual gut. Thus, L. stilbiits must feed often on 0 50 1001s0200 0 50100,s0200 0 50 100110200250300 0 10 100IM200250300 patches of prey. Mean Number per km (S.E.) Yasuda (1960a,b) reasoned that a fish’s gape width determines its ability to trap its prey, while its jaw Figure 19. Diel vertical distribution patterns of Euphausia pacifica, a major prey of S. leucopsarus, in the Santa Barbara (above) and Santa Cruz (below) length determines the size of its prey. Because the basins. All other details as in figure 16. two species have similar gape widths, they should be equally adept at trapping. But S. leucopsavus has the longer jaw and should therefore eat larger prey, both inshore and offshore. In the SBB, 29.5% of the as substantiated by the studies of feeding habits. L. stilbius were parasitized by Aponuviis cnlijofonzicus, In general, the gill rakers of most types of fishes but only 5.7% from the SCB were infected; 4.4% constitute a sieve for filtering and catching food of the S. leucopsums specimens from the SBB were (Martin and Sandercock 1967; Yasuda 1960c; Ya- infected by anisakine nematodes, whereas 7.8% suda and Hiyama 1957). Plankton feeders generally from the SCB were infected. The coefficients of have especially well-developed gill sieves compris- conditions (Cailliet et al. 1986) did not differ be- ing many rakers and accessory processes. Obviously tween species or basin. the rakers of L. stilbius make the more effective bar- rier for retaining smaller prey. The more widely spaced and toothed rakers of S. leucopsunis are prob- DISCUSSION ably better at retaining larger food. Gut length and pyloric caeca may influence size Alimentary Morphology and quantity of food eaten. Groot (1969) and Dar- Alexander (1967) stated that fishes with smaller ne11 (1970) found that fishes with relatively longer mouths can better suck in their prey, whereas fishes guts and many caeca tended to eat smaller prey items with larger mouths can better grasp prey from the and more frequently. In this study L. stilbius, with a side. Therefore, L. stilbius would be better at suck- double stomach, a long intestine, and more caeca, ing in abundant soft items (soft-bodied salps and usually ate large quantities of small, soft foods like larvaceans), and S. leucopsanis would be better at salps, larvaceans, and copepods, whereas S. letico- grasping a greater diversity of larger or more elusive psariis, with its single stomach, short intestine, and prey (copepods and euphausiids). fewer caeca, more often ate single euphausiids and The structure and behavior of soft-bodied, gelat- larger copepods. The rugae and typhlosole ridges inous prey must be considered when interpreting presumably help these fishes process food through how L. rtilbiiis captures them. Larvaceans secrete the cardiac stomachs (Kapoor et al. 1975). houses around themselves, which Alldredge (1976c) All aspects of alimentary morphology, therefore, proposed to be a protective mechanism. Because indicated that L. stilbius is better adapted for contin- L. stilbius appeared to have only the larvacean itself uously gorging itself with abundant smaller and in its gut, either it ingested little of the house, or the often gelatinous prey. In contrast, S. leucopsants house is difficult to detect in stomach contents. Un- should feed more sporadically on smaller quantities derwater observations of L. stilbius indicate that they of larger prey. Indeed, L. stilbius tended to have “re- are relatively passive (Barham 1970). It is possible cently full” stomachs (rather than “recent but not that they can slowly approach larvaceans in their full”), implying that it feeds mostly to fullness. S.

116 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

leticopsavw tended to have more “recent but not full” water (Berner 1957) and presumably not very nutri- stomachs, implying that it feeds more sporadically tious. Optimally, it must eat continuously and and not usually to fullness. digest quickly to meet its energy requirements. L. stilbiiis eats sinall copepods, which may be more Feeding Habits difficult forage, only when the larvaceans and salps Previous studies of food habits of bathylagids are dwindle in numbers. S. leucopsaviis eats a greater size few and sketchy. Hopkins and Torres (1989) found range of more nutritious prey, including large pred- that Buthylugits anturcticcrs ate, among other things, atory crustaceans. gelatinous coelenterates. Anderson (1967) found Comparing the feeding habits of these two fishes that 70% of the cardiac stomachs of L. stilbiits ex- with their growth characteristics produces an appar- amined from the San Pedro Basin contained fish ent paradox. Childress et al. (1980) reported a higher eggs, 60% had copepods, and 44% had fish scales, growth rate for L. stilhius than for S. leircoprunrs, yet while 40% of the pyloric stomachs contained salps, L. stilbiiis consumes prey of relatively lower energy 35% had copepods, 25% had euphausiids, 20% had content. There are three possible explanations of this eggs, aiid 20% had larvaceans. He concluded that paradox. One would be that L. stilhiirs grows large the prey of L. stilbius are less mobile than those of faster, but has tissues that are not as densely con- Tviphotunrs nzexicanur, a common off structed (Childress and Nygaard 1973; Childress et southern California. Noble (1968) found similar al. 1980). A second would be that it expends less prey but noted the dearth of fast, active chaeto- energy than S. leitcopsavirs by not regularly migrat- gnaths in L. stilhitrs stomachs. All studies indicate ing, and by foraging more efficiently on larvaceans that L. stilhius eats relatively small items, although and salps than S. leucoprids does on the larger, more our results showed that larvaceans and ostracods are elusive and vertically migrating crustaceans. A third more abundant food than euphausiids, at least off possibility is that L. stilbiiis is more efficient at as- Santa Barbara. similating the few calories available in its prey. Myctophids in general have been reported to eat copepods, euphausiids, ostracods, mollusks, fish Seasonal Variations in Feeding Habits eggs and larvae, chaetognaths, larval and adult de- The lack of seasonal changes in eating habits and capod shrimp, insects, siphonophores, tunicates, available prey in the relatively eutrophic SBB indi- annelids, sipunculid and nemertine larvae, pycno- cates that food was not limiting there. In contrast, gonids, and foraminifera (Beebe and Vander Pyl the seasonal decrease in food (S. fisfovnzis) available 1944; Aughtry 1953; Paxton 1967b; Anderson 1967; to L. stilbiits in the SCB during the late summer Holton 1969; Legand and Rivaton 1969; Bradbury thermal stratification period may have forced it to and Abbott 1970; Nakamura 1970; Raymont 1970; seek out copepods, which may be harder to catch. Baird et al. 1975a; Gorelova 1975; Hopkins aiid The assumed decrease in primary production dur- Baird 1975; Clarke 1978; Frost and McCrone 1979; ing this period may have caused the coincident de- Kinzer and Schultz 1985; Young and Blaber 1986; cline in salp (and presumably larvacean) catches in Dalpadado and Gjosaeter 1988). the trawls, as seen by Hubbard and Pearcy (1 971) off Our results gerierally correspond with previous Oregon. These filter-feeding organisms require studies, which found that euphausiids and calanoid high concentrations of phytoplankton, and thus copepods constitute most of the S. leucopsuvzis diet flourish in replenished surface waters enriched by (Bary et al. 1962; Osterberg et al. 1964; Paxton nutrients brought up from unstratified depths (Sil- 1967b; Tyler and Pearcy 1975; Collard 1970). How- ver 1975). ever, fish from the SBB ate the shrimp Seyestes Also, in the SBB L. stilhitis had few potential siniilis much less frequently than did fish from competitors, and offshore in the SCB there were Monterey Bay (Barhatn 1957), and contained fewer only a few more. Its more oceanic relative Butlzylups amphipods than fish from Saanich Inlet, British Co- wescthi eats larvaceans and salps (M. Kelley, pers. lumbia (Bary et al. 1962). No other studies reported coinin.), but almost never enters inshore waters, and as high a frequency of ostracods as we found in invades the SCB in noticeable numbers only during SBB fish. the fall thermal stratification period. Even then it is L. stilhiits and S. leiicop~uurshave very different far less abundant than L. stilhiiis (cf. Brown 1974). feeding habits. L. stilbirtr ingests a relatively narrow Farther offshore, however, Bathyhgus spp. far out- variety of prey. It eats large amounts of small, slug- number L. stilhirts (A. Ebeling, unpublished data). gish, herbivorous jellies, which are 90% -95% In the SCB, Buthylups spp. may compete only dur-

117 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

ing the warming season, when, coincidentally, retreat to deeper waters during the daytime to rest, L. stilbius ate more copepods and fewer jellies. digest, and avoid predation (cf. McLaren 1963; Pax- Offshore, S. leucopsavus is probably more abun- ton 1967a; Nafpaktitis 1968; Marshall 1954, 1980). dant than L. stilbiiis because it is a generalized pred- Indeed, the common prey of both fishes tend to in- ator andemay broaden or narrow its diet as the habit the upper 400 m, and many of them migrate situation demands. Its feeding habits are much more vertically in a diel pattern. similar between basins than those of L. stilbius, Several authors have observed that L. stilbius con- which may have to broaden its diet beyond optimal centrate at mid-depth during daytime but broaden limits in deeper waters offshore, where its preferred their vertical distribution by dispersing upward at salps and larvaceans are not so concentrated and night, usually not in a distinct layer (Anderson 1967; evenly distributed among seasons. Although Tucker 1951; Clarke 1970; Ebeling et al. 1970b). S. leucopsavus can eat many different items both in- Other authors contend, from direct observations shore and offshore, it can also feed on either euphau- made off San Diego from deep submersibles, that siids or copepods, depending on how the food L. stilbius seldom ascend above 500 m and therefore supply changes. Because S. leiicopsavus can eat just do not exhibit a daily migratory pattern, but at times about anything it encounters, competition for items they do come to the surface in large numbers (Bar- like copepods and ostracods may be less in the SBB. ham 1970; Pickwell et al. 1970). This somewhat The tendency, during seasons I1 and 111, for off- unpredictable behavior helps explain the high vari- shore S. leucopsavus to eat mostly euphausiids, also ability in the vertical distribution data, especially in observed by Collard (1970), could be explained by the SBB (figure 12). competition or prey availability. During these pe- To optimize its feeding strategy, L. stilbius must riods, myctophid competitors belonging to an “off- sometimes visit the surface waters where larvaceans shore fish group” (cf Ebeling et al. 1970a) become and salps occur. Our samples indicated that most of seasonally abundant and may force S. leucopsanis to these fish descended before daylight, although pos- restrict its diet. An alternate explanation is that eu- sible laggards may avoid the trawl in sunlit waters phausiids may become more abundant. S. leucopsaviis during the daytime. Our medium-speed trawls may may broaden its diet to include more copepods dur- have caught them effectively in the dark but not ing the cold winter season when the offshore fishes during the daytime. But occasionally our trawl did dwindle in numbers. catch many individuals near the surface during the day. Unfortunately, few shallow hauls were made Diel Vertical Migrations during the day in the SCB. Even though fish abundances were standardized Many authors have noted the diel vertical migra- by trawling effort, abundances varied considerably tion of S. leiicopsavus; the fish is one component of among collections. For either species, this variability the sonic scattering layer and tends to respond to a could be a function of disjunct distributions among specific isolume (Tucker 1951; Barham 1957; Fast depth zones, areas, or seasons; differences in ability 1960; Bary et al. 1962; Pearcy and Laurs 1966; Pax- to avoid the net, which is size-specific for fishes ton 1967a; Taylor 1968; Bary and Pieper 1970; Bar- (Aron and Collard 1968); or a tendency to occur in ham 1970; Clarke 1970; Pearcy and Mesecar 1970; clumps (Pearcy 1964; McGowan and Fraundorf Ebeling et al. 1970a,b). Others have also noted that 1966; Harrisson 1967; Alldredge et al. 1984). Avoid- not all of the population ascend toward the surface ance or escape may be more important during the waters every night (Paxton 1967b; Barham 1970; day than at night (Pearcy and Laurs 1966). However, Clarke 1970; Zahuranec and Pugh 1970). any daytime avoidance could be negated if either species is lethargic at diurnal depths, as indicated by Fullness and Recency of Feeding Relative to Barham (1970), or it may be enhanced if they are Vertical Migration hanging there but are quite ready to flee at the Prey of different species are most likely digested approach of a predator or midwater trawl (cf. Rob- at varying rates under different conditions (Windell ison 1972). 1967). In general, the stomachs of small fishes prob- Since surface waters contain more food than deep ably empty in about 12 hours (Anderson 1967; Tyler waters (Vinogradov 1974; Marshall 1954, 1980), 1970). However, since all four categories of fullness L. stilbiiis and S. leucopsanis, like many other meso- and recency of feeding occurred in both species, it pelagic fishes, should benefit from regular feeding should not matter how long digestion takes because migrations toward the surface at night. They may the recency and fullness indices will be relative. Es-

118 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

timates of how recently a particular gut was filled, night, where it can use its large eyes to find salps and however, are not possible without data on digestion larvaceans, and be protected from predation by its rates (Hopkins and Baird 1977). silvery coloration. The rest of the time it can find The digestibility of the common prey of L. stilbius refuge from surface predators in deeper waters. S. may shed some light on the fullnesslrecency data and leucopsavus, on the other hand, with its the feeding cycle of this species. Shelbourne (1962) and large mouth, most likely migrates to deeper observed that soft tissues of Oikopleuva were quickly water to seek refuge from predation, but can feed at digested after capture by larval plaice. This implies all times and depths on copepods and co-migrating that the mostly intact larvaceans in recently full euphausiids. Thus its regular migration to the sur- stomachs of L. stilbitis were newly ingested. In the face it not obligatory, because the fish can consume only other study of the feeding cycle of this species, prey at any depth. Yet vertical migrations may place Anderson (1967) found that the guts of L. stilbitis in these fishes in surface currents that might help them the Catalina Basin were fuller with only partially find concentrations ofprey (Isaacs et al. 1974). digested material at night than during the day. He suggested that L. stilbius feeds readily at the surface Factors Controlling Abundance of L. stilbius and at night, but he could not determine if there was S. leucopsarus much feeding at greater depths. Our relatively high Several possible factors may explain why L. stil- percentages of “recently full” stomachs, as com- bius is less successful offshore than S. leucopsanis. pared with “recent but not full” stomachs among Both species appear to breed successfully in both fish caught in the surface waters indicate that places. Analysis of egg sizes versus size of females L. stilbius feeds to fullness whenever possible. (Childress et al. 1980) indicates that both species Existing studies of S. leucopsavus compare favor- attain sexual maturity in the two localities. Also, ably with ours in that the fish were found to feed larvae of the two fishes occur abundantly in both mostly at night near the surface (Anderson 1967; basins (Ahlstrom 1965). In the SBB, 50% of all Holton 1969). But they were also found to feed in L. stilhius captures and ’45% of all S. leucopsavus cap- the morning and afternoon (Paxton 1967b; Tyler and tures were of young (<50 mm SL) or larvae; in the Pearcy 1975). However, the digestibility of prey SCB, the figures were 70% for L. stilbius and 50% consumed by S. leucopsanis must be interpreted dif- for S. leucopsarus (Brown 1974). Ebeling et al. ferently. Since this fish eats mostly crustaceans, (1970b) also concluded that all growth stages of digestion may take several hours. Therefore, rela- these two common fishes were abundant in both tively undigested items may persist in the stomachs places. of deep fish that had fed earlier in shallower waters. Even though the two species differ in seasonal Like L. stilbius, S. leticopsavus never had a high per- abundances, there is no evidence that they enter or centage of “full but not recent” stomachs, implying leave the two basins at different rates. Brown (1974) that the fish clear their stomachs rapidly. Because reported that the more physiographically and many of their stomachs were empty, and because hydrographically isolated inshore SBB restricted their proportion of “recent but not full” stomachs faunal intrusions from the offshore oceanic environ- often exceeded their proportion of “full and recent” ment. Ebeling et al. (1970a) defined an offshore fish stomachs for most depths, these fish may feed group consisting of “tropical” species, which in- whenever they can, mostly on larger, less digestible crease in numbers in the SCB and farther offshore items. Thus they seldom completely fill their stom- during the summer and fall when the California achs. Also, the primary prey of S. leticops?stlnis are Current weakens. L. stilbius and S. leucopsarus co-oc- found between 0 and 200 m at night and between cur with these “offshore fishes, ” but both species 200 and 400 m during the day (Viriogradov 1968, also belong to a resident “inshore” community of 1974; Youngbluth 1976; figure 19). The ultimate midwater animals, which abound in both basins resolution of this question awaits an evaluation of throughout the year. Their otoliths, found in bot- digestion rates of mesopelagic fishes at different tom cores (Soutar and Isaacs 1969), indicate that temperatures (Gorelova 1975; Hopkins and Baird both species have occupied the SBB for at least 2,000 1977; Young and Blaber 1986; Dalpadado and Gjo- years. saeter 1988; Kinser and Schultz 1985). Despite differences in their parasite infection These two midwater fishes appear to benefit from rates, both species seem equally healthy and robust vertical migration in different ways. L. stilbius can in the two basins. L. stilhitts is more heavily parasi- occupy surface waters, either in the afternoon or at tized by trematodes in the SBB than the SCB (Noble

119 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

1968; Noble and Orias 1970). S. leucopsuvus, on the hand, because salps and larvaceans filter out small other hand, is more heavily parasitized by cestodes particles in the water column, including phyto- in the SBB but is equally infected with nematodes plankton (Silver 1975; Alldredge 1976a), and are in in both basins (Noble and Collard 1970). Our obser- turn consumed by other micronekton (Alldredge vations of parasitism concur with the above studies, 1976b; Michaels and Silver 1988), they must provide and condition factors did not significantly differ energy throughout the open water column (Morris within species between basins. et al. 1988). Indeed, L. stilbitis and S. leucopsuvus, Predation does not seem likely to be a key factor. through their consumption of salps, larvaceans, and The same kinds of predators, with the possible ex- crustaceans, and through their vertical migrations, ception of relatively large, deep-sea fishes such as must play an active role in transporting energy Chutrliodus (Borodulina 1973), occur in both places sources from the surface to deeper water (cf. Pearcy and could eat both species. It is possible that the et al. 1977), at least in the form of fecal matter, which more bathymetrically compressed SBB habitat sinks at several cm sec-' (Robison and Bailey 1981). could concentrate more predators, but there is no Research is needed to determine if these fishes influ- evidence of this. ence the rates of vertical flux of organic matter in the Deepsea smelts are reportedly eaten by rockfish open ocean. (Lambert 1960), albacore (McHugh 1952), and ce- taceans (Fitch and Brownell 1969). In particular, ACKNOWLEDGMENTS L. stilbius is eaten by albacore and bluefin tuna (Pin- We thank R. W. Holmes, J. H. Connell, E. R. kas et al. 1971). One Chiasmodon niger, a predatory Noble, B. Robison, and B. Nafpaktitis for critically bathypelagic fish from the San Clemente Basin, had reviewing the original manuscript. G. S. Arita, D. a large adult L. stilbitrs in its distended stomach (Bor- W. Brown, S. B. Collard, E A. DeWitt, Jr., H. odulina 1973). Genthe, L. Hendrian, R. M. Ibara, M. S. Love, €? Likewise, myctophids are reportedly eaten by a Setzer, and H. Tyler helped with the field work and variety of predators, including cephalopods; large laboratory data analyses. R. M. Ibara and M. Palm- pelagic fishes such as sharks, , rockfishes, and gren wrote computer programs that helped with the swordfish; other deep-sea fishes; sea birds; and ma- statistical analyses. The University of California, rine mammals (e.g., Marshall 1954; Paxton 1967b; Santa Barbara Computer Center provided addi- Tyler and Pearcy 1975; Ainley et al. 1986). In partic- tional computer time. This study was supported by ular, S. leucupsuvtrs has been eaten by sharks (Hubbs NSF grants GB 2867 and 4698 for ship time and GB 1917), salmon (Shimada 1948), albacore and bluefin 4669 and 7973 for the shore-based analyses. A Uni- tuna (Pinkas et al. 1971), rockfishes (Eigenmann and versity Fellowship, University of California, Santa Eigenmann 1890; Starks and Morris 1907; Pereyra Barbara, provided the senior author with one year et al. 1969), and cetaceans (Fitch and Brownell 1969). of support. It therefore seems most probable that feeding habits best account for the differential success of LITERATURE CITED these two mesopelagic fishes offshore. The way in Ahlstrom, E. H. 1965. Evaluation offishes resources ofthe Pt. Arguello which they use the available food resources may area. Part I. Fish larvae of the Pt. Arguello area. Report AT (49-7)- have a great deal to do with their relative success in 2428, Division of Biology and Medicine, U.S. Atomic Energy Commission. different habitats. -. 1969. Mesopelagic and bathypelagic fishes in the California Current Region. Calif. Coop. Oceanic Fish. Invest. Rep. 13: 39-44. Potential Role of These Fishes in Energy Transport Ainley, D. G., W. R. Fraser, C. W, Sullivan, J. J. Torres, T. L. Hopkins, and W. 0. Smith. 1986. Antarctic mesopelagic micronekton: evi- to the Deep Sea dence from sea birds that pack ice affects community structure. Sci- In spite of their apparent lack of nutritional value, ence 232: 847-849. numerous midwater fishes have been reported to Alexander, R. M. 1967. Functional design in fi5hes. London: Hutchin- SO11 311d CO., 164 pp. feed on salps and larvaceans (e.g., Gorelova 1974, Alldredge, A. L. 1967a. Appendicularians. Sci. Am. 235(1):94-102. 1975; Baird et al. 1975b; Kashkina 1986; Longhurst -. 1976b. Discarded appendicularian houses as sources of food, and Harrison 1988; Hopkins and Torres 1989). How surface habitats, and particulate organic matter in planktonic environ- ments. Limnol. Oceanogr. 21(1):14-23. these fishes utilize the gelatinous zooplankters is still -. 1976c. Field behavior and adaptive strategies of appendiculari- uncertain. Kashkina (1986) proposed that the tunica ails (Chordata: tunicata). Mar. Biol. 38:29-39. is only partially assimiliated, if at all, and it must Alldredge, A. L., B. H. Robison, A. Fleminger, J. J. Torres, J. M. King, and W. M. Hamner. 1984. Direct sampling and in situ observation of take considerable energy for a fish to consume suf- a persistent copepod aggregation in the mesopelagic zone of the Santa ficient material to constitute a meal. On the other Barbara Basin. Mar. Biol. 80:75-81.

120 CAlLLlET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES CalCOFl Rep., Vol. 31,1990

Anderson, R. 1967. Feeding chronology in two deep-sea fishes off Cal- Patterns of growth, energy utilization and reproduction in some ifornia. M. S. thesis, University of Southern California. meso- and bathypelagic fishes off southern California. Mar. Biol. Aron, W. I., and S. B. Collard. 1968. A study of the influence of net 61 27-40, speed on catch. Limnol. Oceanogr. 14242-249. Clarke, T. A. 1978. Diel feeding patterns of 16 species of niesopelagic Aron, W. I., N. Raxter, R. Noel, and W. Andrews. 1964. A description fishes from Hawaiian waters. Fish. Bull. 76(3):495-513. of a discrete depth plankton sampler with some notes on the towing Clarke, W.D. 1970. Comparison of different investigative techniques behavior of a 6-foot Isaacs-Kidd midwater trawl and a 1-m ring net. for studying the deep scattering layers. 111 Proc. Internat. Synip. Biol. Limnol. Oceanogr. 9:324-333. Sound Scattering in the Ocean, G. B. Farquhar, ed. Maury Center Aughtry, R. H. 1953. A note on the mass mortality of the myctophid Ocean Sci., Washington, D.C. No. 5, pp. 551-562. fish TavletoriLcnrria rveriitlaris. Copeia 1953(3):190-192. Collard, S. B. 1970. Forage of some eastern Pacific midwater fishes. Baird, R. C., T. L. Hopkitis, and D. F. Wilson. 1975a. Diet and feeding Copeia 1970(2):348-354. chronology of Dioplzits raariirigi (Mytophidae) in the Cariaco Trench. Dalpadado, I?, and J. Gjosaeter. 1988. Feeding ecology ofthe lanternfish Copeia 1975(2):356-365. Berirlzoscrrraprevotttm from the Indian Ocean. Mar. Biol. 99:555-567. Baird, R. C., N. €? Thompson, T. L. Hopkitis, and W. R. Weiss. 1975b. Darnell, R. M. 1970. Evolution aiid the ecosystem. Am. Zool. Chlorinated hydrocarbons in mesopelagic fishes of the eastern Gulf 10:9-17. ofMexico. Bull. Mar. Sci. 25(4):473-481. Ebeling, A. W., G. M. Cailliet, R. M. Ibara, E A. DeWitt, Jr.. and D. Barham, E. G. 1957. The ecology of sonic scattering layers in the Mon- W. Brown. 1970a. 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Deep-sea Res. 13:153-165. Lavenberg, R. J., and A. W. Ebeling. 1967. Distribution of midwater Pearcy, W, G., and R. S. Mesecar. 1970. Scattering layers and vertical fishes among deep-water basins of the southern California shelf. In distribution of oceanic animals off Oregon. IIZProc. Internat. Synip. Proc. Symp. Biol. Calif. Islands, Santa Barbara Botanic Garden, Biol. Sound Scattering in the Ocean, G. B. Farquhar, ed. Maury R. M. Philbrick, ed. Pp. 185-201. Center Ocean Sci., Washington, D.C. No. 5, pp. 381-394. Legand, M., and J. Rivaton. 1969. Cycles biologiques des poissons Pearcy, W. G., E. E. Krygier, and N. H. Cutshall. 1977. Biolog- mesopelagiques de l’est de l’Ocean Indien. Troisieme note: action ical transport of zinc-65 into the deep sea. Limnol. Ocemogr. predatrice des poissons micronectoniques. Cah. ORSTOM (ser. 22(5):846-855. Oceanogr.) 7:29-45. Pereyra, W. T., W. G. Pearcy, and F. E. Carvey, Jr. 1969. Srbastodes Longhurst, A. R. 1967. Diversity and trophic structure of zooplankton javidns, a shelf rockfish feeding on mesopelagic fauna, with con- communities in the California Current. Deep-sea Res. 14:393-408. sideration of the ecological implications. J. Fish. Res. Bd. Can. Longhurst, A. R., and W. G. Harrison. 1988. Vertical nitrogen flux 25:221 1-221 5. from the oceanic photic zone by diel migrant zooplankton and nek- Pickwell, G. V., R. J. Vent, E. G. Barham, W. E. Batzler, and I. E. ton. Deep-sea Res. 35(6):881-889. Davies. 1970. Biological acoustical scattering off southern California, MacArthur, R. H., and J. H., Connell. 1966. The biology of popula- Baja California, and Guadalupe Island. Iiz Proc. Internat. Synip. Biol. tions. New York: John Wiley, 200 pp. Sound Scattering in the Ocean, G. B. Farquhar, ed. Maury Center Malone, T. C. 1971. The relative importance of nannoplankton and Ocean Sci., Washington, D.C. No. 5, pp. 490-507. netplankton as primary producers in the California Current system. Pinkas, L., M. S. Oliphant, and I. L. K. Iverson. 1971. Food habits of Fish. Bull. 69:799-820. albacore, bluefin tuna, and bonito in California waters. Calif. Fish Marshall, N. B. 1954. Aspects of deep-sea biology. London: Hutchin- Game, Fish Bull. 152:l-105. son, 379 pp. Raymont, J. E. G. 1970. Problems of the feeding of zooplankton in the -. 1980. Deep-sea biology: developments and perspectives. New deep sea. In Proc. Internat. Symp. Biol. Sound Scattering in the York: Garland STPM Press, 566 pp. Ocean, G. B. Farquhar, ed. Maury Center Ocean. Sci., Washington, Martin, N. V., and F. K. Sandercock. 1967. Pyloric caeca and gill raker D.C. No. 5, pp. 136-148. development in lake trout, Saltdinus ~amaycush,in Algonquin Park, Robison, B. H. 1972. Distribution of the midwater fishes of the Gulf of Ontario. J. Fish. Res. Bd. Can. 24:965-974. California. Copeia 1972(3):448-461. McAllister, D. E. 1961. A collection of oceanic fishes from off British Robison, B. H., and T. G. Bailey. 1981. Sinking rates and dissolution Columbia with a discussion of the evolution of black peritoneum. of midwater fish fecal matter. Mar. Biol. 65:135-142. Bull. Nat. Mus. Can. 17239-43. Shelbourne, J. E. 1962. A predator-prey size relationship for plaice lar- McGowan, J. A,, and V. J. Fraundorf. 1966. The relationship between vae feeding on 0ikopIerrr.a.J. Mar. Biol. Assn. U.K. 42243-252. size of net used and estimates of zooplankton diversity. Limnol. Shimada, B. M. 1948. Records of lanternfish in Puget Sound. Copeia Oceanogr. 11:456-469. 1948(3):227. McHugh, J. L. 1952. The food of albacore (Germo alalurrga) off Califor- Sholkovitz, E. R., and J. M. Gieskes. 1971. A physical-chemical study nia and Baja California. Bull. Univ. Calif. Scripps Inst. Oceanogr. of the flushing of the Santa Barbara Basin. Limnol. Oceanogr. 6: 161-1 72. 16:479-489. McLaren, I. A. 1963. Effects oftemperature on growth of zooplankton, Silver, M. 1975. The habitat of Salpa fusfefor.mis in the California and the adaptive value of vertical migration. J. Fish. Res. Bd. Can. Current as defined by indicator assemblages. Limnol. Oceanogr. 20:685-727. 20(2) 230-237. Michaels, A. F., and M. W. Silver. 1988. Primary production, sinking Sokal, R. R., and F. J. Rohlf 1969. Biometry. San Francisco: W. H. fluxes and the microbial food web. Deep-sea Res. 35(4):473-490. Freeman and Co. 776 pp. Miller, D. J., and R. N. Lea. 1972. Guide to the coastal marine fishes of Soutar, A., and J. D. Isaacs. 1969. History of fish populations inferred California. Calif Fish Game, Fish Bull. 157:l-235. from fish scales in anaerobic sediments off California. Calif. Coop. Morris, R. J., Q. Bone, R. Head, J. C. Braconnot, and l? Nival. 1988. Oceanic Fish. Invest. Rep. 13:63-70. Role of salps in the flux of organic matter to the bottom of the Lig- Starks, E. C., and E. L. Morris. 1907. The marine fishes of southern urian Sea. Mar. Biol. 97(2):237-242. California. Univ. Calif Publ. Zool. 3:159-251. Nafpaktitis, B. G. 1968. Taxonomy and distribution of the lantern- Taylor, E H. C. 1968. The relationship of midwater trawl catches to fishes, genera Lobianchia and Diaphus, in the North Atlantic. Dana- sound scattering layers off the coast of northern British Columbia. Rept. 73:l-131. J. Fish. Res. Bd. Can. 25:457-472. Nakamura, E. L. 1970. Observations on the biology of the myctophid Tucker, G. 1951. Relations of fishes and other organisms to the scatter- Diaphusgarmani. Copeia 1970(2):374-377. ing ofunderwater sound. J. Mar. Res. 10215-238. Noble, E. R. 1968. The flagellate Cryptobia in two species of deep sea Tyler, A. V. 1970. Rates ofgastric emptying in young cod. J. Fish. Res. fishes from the eastern Pacific. J. Parasit. 54:720-724. Bd. Can. 27:1177-1189. Noble, E. R., and S. B. Collard. 1970. The parasites ofmidwater fishes. Tyler, H. R., and W. G. Pearcy. 1975. The feeding habits ofthree species Iii A symposium on diseases of fishes and shellfishes, S. Sniesko, ed. of lanternfishes (Family myctophidae) off Oregon, USA. Mar. Biol. Spec. Publ. Am. Fish. Soc. No. 5, p. 57068. 32:7-11.

122 CAILLIET AND EBELING: FEEDING OF TWO VERTICALLY MIGRATING MESOPELAGIC FISHES ColCOFl Rep., Vol. 31,1990

Vinogradov, M. E. 1968. Vertical distribution of the oceanic zooplank- -. 1960c. The relationship of the gill structure and food habits of ton. Israel Program for Sci. Transl., Jerusalem (1970), 330 pp. some coastal fishes in Japan. Rec. Oceanogr. Works Japan 5:139-152. -. 1974. Depth of the nighttime rise of deep scattering layers in Yasuda, F., and Y. Hiyama. 1957. Mechanism of utilization of plankton the central Pacific. Oceanology 14(6):891-895 (Translated by Scripta by some fishes. Rec. Oceanogr. Works Japan 3:85-91. Technica, Inc. for the Am. Geophys. Union). Young, J:W., and S. J. M. Blaber. 1986. Feeding ecology ofthree species Willis, J. M., and W. G. Pearcy. 1980. Spatial and temporal variations in of midwater fishes associated with the continental slope of eastern the population size structure of three lanternfishes (Myctophidae) off Tasmania, Australia. Mar. Biol. 93:147-156. Oregon, USA. Mar. Biol. 57:181-191. Youngbluth, M. J. 1976. Vertical distribution and diel migration of Windell, J. T. 1967. Rates of digestion in fishes. In The biological basis euphausiids in the central region ofthe California Current. Fish. Bull. offreshwater fish production, S. Gerking, ed. New York: John Wiley, 74(4):925-936. pp. 151-173. Zahuranec, B. J,, and W. L. Pugh. 1970. Biological results from scatter- Yasuda, E 1960a. The feeding mechanism in young fishes. Rec. Ocean- ing layer investigations in the Norwegian Sea. It1 Proc. Internat. ogr. Works Japan 5:132-138. Symp. Biol. Sound Scattering in the Ocean, G. B. Farquhar, ed. -, 1960b. The feeding mechanisms in some carnivorous fishes. Maury Center Ocean Sci., Washington, D.C. No. 5, pp. 360-380. Rec. Oceanogr. Works Japan 5:153-160.

123 THOMSON: U.S. MARKET FOR FISH MEAL AND OIL CalCOFl Rep., Vol. 31,1990

THE MARKET FOR FISH MEAL AND OIL IN THE UNITED STATES: 1960-1988 AND FUTURE PROSPECTS

CYNTHIA J. THOMSON National Marine Fisheries Service Southwest Fisheries Center P. 0.Box 271 La Jolla, California 92038

ABSTRACT queria del bacalao de Alaska, (4) 10s esfuerzos para Fish meal is used in the United States largely as a desarrollar productos de consumo direct0 por el ser high-protein ingredient in poultry feed. Prices of humano, y por ultimo (5) el estado de las pesquerias domestic fish meal are determined by world market en el Jap6n y en SudamCrica. conditions for fish meal as well as other oil meals. Faced with limited fish meal supplies and little con- INTRODUCTION trol over prices, the U. s. poultry industry has sub- The fish meal and oil industry began in the nine- stituted other ingredients and made use of tech- teenth century in northern Europe and North nological advances to satisfy the nation’s growing America. The oil was manufactured from surplus demand for table birds. fish caught in the herring fishery, and was used to A number of factors have been identified that may tan leather and as an ingredient in products such as significantly affect future demand, supply, and soap. The residue was originally used as fertilizer. prices of fish meal and oil, both in the United States However, since the turn of this century, the use of and abroad. These include (1) increases in world fish for fertilizer has diminished considerably be- aquaculture production, (2) possible development of cause increasing amounts have been diverted to the a domestic market for hydrogenated fish oil, production of fish meal (FA0 1986). (3) recent changes in the Alaska pollock fishery, The fish meal production process, which is (4) efforts to develop marketable products for direct known as reduction, involves cooking the fish, re- human consumption from reduction species, and moving the water and oil from it, drying the solid (5) the status of the Japanese and South American material left behind, and grinding it into a meal. Fish sardine fisheries. meal is used in the United States largely as a high- protein ingredient in poultry feed. It is also used in RESUMEN feeds for pigs, farmed fish, fur-producing animals, La harina de pescado es un ingrediente de alto laboratory animals, and household pets. contenido proteico utilizado corrientemente en la The oil that is removed in the reduction process is alimentacion de aves de corral en 10s Estados Unidos marketed as an ingredient for industrial products de NorteamCrica. El mercado mundial determina el (such as paints and lubricants) and foods (such as precio de la harina, como asi tambitn el de otras margarines and shortenings). Most of the fish oil harinas de orujo. Ante el problema de un limitado produced in the United States has historically been abastecimiento de harina de pescado y un reducido exported to Europe for use in margarines. Until control en 10s precios, la industria de la polleria se very recently, the U. S. Food and Drug Administra- vi6 obligada a sustituir otros ingredientes y hacer tion did not allow the use of fish oil in products for us0 de 10s avances tecnologicos con el fin de satis- domestic human consumption. facer la creciente demanda de estas aves de corral. Approximately one metric ton (MT) of fish meal Se han identificado un nfimero de factores que is produced from each 4-5 MT of fish harvested. pueden afectar significativamente la futura de- The oil yield is more variable and depends on the manda, el abastecimiento y el precio de la harina de species and the time of year when the fish are caught pescado y otros aceites, tanto en 10s Estados Unidos (Vondruska 1980). For example, the oil content of como en el exterior, a saber: (1) el aumento en la northern anchovy is low during the winter and produccidn de las piscifactorias, (2) el posible desa- spring spawning period, and highest in late summer rrollo de un mercado domestico de aceite de pescado (Lasker and Smith 1977). The wholesale value of hidrogenado, (3) 10s cambios recientes en la pes- U. S. fish meal production has historically exceeded the value of oil production by a factor of two to four ~ [Manuscript received February3,1990 ] (table 1).

124 THOMSON: U.S. MARKET FOR FISH MEAL AND OIL ColCOFl Rep., Vol. 31,1990

TABLE 1 substantial reduction fishery once existed in Califor- Quantity and Value of Fish Meal and Oil Produced in the nia for Pacific sardine (Suvdinops sapx), but the fish- United States ery collapsed in the early 1950s (Radovich 1981). The Production quantity Value state of California lifted its moratorium on sardine (1000's of metric tons) (millions of dollars)

~~ ~~ ~ ~~ ~~~ ~~~ landings in 1986, but so far has allowed modest har- Year Fish meal Fish oil Fish meal Fish oil vests to be taken for nonreduction uses only. - -~ ~- ~~ - ~~ 1960 263.3- 94.8 25.3 13.0 Although most fish meal is produced from whole 1961 282.4 117.1 31.9 14.3 fish, about 10% is produced from the by-catches and 1962 283.3 113.4 35.6 11 .o 1963 -137 7 84.3 30.2 10.9 byproducts of other fisheries. Examples are tuna- 1964 213.5 81.7 28.0 13.3 mackerel and pollock meals, which are produced 1965 230.5 88.7 35.7 14.9 from the scraps remaining after these species are 1966 203.4 74.7 32.3 12.5 1967 191.6 55.5 26.0 6.1 processed into other market products. 1968 213.3 79.0 30.3 7.3 As indicated in table 3, U. S. fish meal production 1969 229.2 77.0 39.8 9.3 ranged from 200 to 350 thousand MT per year from 1970 241.2 93.5 46.4 18.2 1971 265.6 120.4 41.5 20.8 1960 through 1988. Before 1982, menhaden meal 1972 259.0 85.5 48.3 13.1 constituted 55%-80% of annual production. Since 1973 253.2 101.9 119.1 2.5.6 1982, menhaden's share has been even higher, aver- 1974 264. 6 107.9 83.5 49.2 1975 253.4 111.4 64.6 32.6 aging 85% of total production. Tuna-mackerel meal 1976 271.3 92.8 95.7 31.2 contributes 20,000-45,000 MT per year. 1977 248.4 60.7 96.5 28.4 Anchovy meal production peaked in 1975 at 1978 320.9 134.4 120.2 60.7 1979 329.2 121.4 133.3 i4.1 25,100 MT, when its share of total production was 1980 322.3 141.8 132.9 57.9 10%. However, meal production from this species 1981 281.3 83.0 117.6 33.1 has been much lower in most other years and has 1982 330.4 157.6 121.2 53.6 1983 339.0 181.1 129.1 66.8 been negligible since 1983, largely because of eco- 1984 334.7 169.1 112.6 61.0 nomic factors rather than low abundance. From 1983 1985 319.6 120.3 83.1 41 .9 through 1988 the fish meal price ranged from $240 1986 308.1 152.8 82.4 43.7 1987 349.6 135.4 120.9 35.5 to $440 per MT, and the ex-vessel price received by 1988 3-83.5 101.9 129.2 43.6 the menhaden fleet ranged from $80 to $115 per MT.

~~ ~ ~~ References: U. S. Department of the Interior 1960-1970; U. S. Depart- However, for reasons that are not clear, the ex-vessel ment ofCommercc 1971-1988. anchovy price offered by California processors has remained at record low levels (below $35 per MT). As a result, the California reduction fleet has not Fish and shellfish landed commercially in the found it profitable to target on anchovy and has di- United States are used for human consumption and rected increasing amounts of effort to more lucrative industrial products. Over 85% of industrial use is species such as mackerel, tuna, and squid (Thomson attributable to reduction; the remainder consists of et al. 1989). bait and animal food. Since 1960, industrial uses have accounted for 36%-53% of total landings but only 4°/o-110/~ of total ex-vessel revenues on an an- Imports and Exports nual basis (table 2). Reduction landings tend to be Figure 1 describes the contributions of Peru and large in quantity but low in value relative to landings Chile to world exports of fish meal. From 1960 to used for direct human consumption. 1972 Peruvian anchoveta (Engratrlis rirz'qevzs) ac- counted for 50%-63% of all the fish meal traded in international markets. A combination ofoverfishing SUPPLY and poor recruitment led to the collapse of the fish- ery during the 1972-73 El Nifio (Glantz 1979). The Production recovery of the fishery has enabled Peru to signifi- The species used for reduction are small, oily, pe- cantly increase its exports in the 1980s, though not lagic fishes that are not marketable in large quan- to the high levels of the 1960s and early 1970s. Chile's tities for human Consumption. In the United States, fish meal exports began to increase in the early theseinclude northern anchovy (Engruttlis mordax) on 1970s, as a result of its developing sardine (Sar.dirzc7ps the Pacific Coast and menhaden (Buevoortiu tyiwvznus sqpx) and jack mackerel (E.ncCzwirs mtryhyi) fisher- and B. patronus) on the Atlantic and Gulf coasts. A ies. Since 1980 Chile's exports have exceeded Peru's.

125 THOMSON: US. MARKET FOR FISH MEAL AND OIL CalCOFl Rep., Vol. 31,1990

TABLE 2 U.S. Commercial Finfish and Shellfish Landings and Ex-Vessel Value (Millions of Dollars) by Disposition of Catch Human food Industrial useb Total __ ~ .~ ~~~ ~~~ ~~~~~~~~ ~ Year Landings' Value Landings Value Landings Value

~~~~~~~~~~~~ ~~~ ~~ ~~ ~~ ~- .. .. ~ ...~ ~~ ~ ~~ 1960 1133.1 - 1108.6 - 2241.7 354 1961 1129.5 - 1223.4 - 2352.8 362 1962 1152.1 - 1276.4 - 2428.6 396 1963 1159.4 - 1039.2 - 2198.6 377 1964 1132.6 - 927.2 - 2059.8 389 1965 1173.6 408 993.2 29 2166.8 446 1966 1166.9 437 813.5 24 1980.4 472 1967 1073.9 414 765.3 17 1839.1 440 1968 1064.1 468 822.8 29 1887.0 497 1969 1052.8 492 91 4.5 35 1967.3 527 1970 1150.8 565 1079.6 48 2230.3 613 1971 1107.2 604 1168.9 47 2276.1 65 1 1972 1104.5 702 1075.3 46 2180.0 748 1973 1087.7 836 1115.8 101 2203.6 937 1974 1132.2 844 1120.8 88 2253.0 932 1975 1118.1 904 1094.1 73 2212.2 977 1976 1258.7 1257 1185.2 92 2444.0 1349 1977 1339.0 1440 1051.9 114 2390.9 1554 1978 1441.1 1733 1293.2 121 2734.3 1854 1979 1505.0 2093 1337.7 141 2842.7 2234 1980 1657.4 2092 1282.8 145 2940.2 2237 1981 1608.9 2277 1102.2 111 2711.1 2388 1982 1490.1 2247 1398.0 143 2888.1 2390 1983 1468.7 2203 1452.0 152 2920.7 2355 1984 1505.9 2206 1414.3 144 2920.3 2350 1985 1494.1 2198 1344.5 128 2838.6 2326 1986 1539.1 2641 1196.6 122 2735.6 2763 1987 1789.9 2979 1338.1 136 3128.0 3115 1988 2081.1 3362 1181.2 158 3262.3 3520 .. ~~~~~ ~ ~ ~ ~~ ~~~ ~~~ 'Thousands of metric tons, round weight; excludes weight of mollusk shells. I, Over 85% processed into meal, oil, and solubles. The remainder is used for shell products, bait, and animal food. References: U. S. Department of the Interior 1960-1970; U. S. Department of Commerce 1971-1988.

The two countries together account for 40% -50% imported supplies from 1960 through 1972. Peru- of world fish meal exports in the 1980s. vian imports declined significantly after the collapse U.S. imports of fish meal have tended to follow of the anchoveta fishery, and the resulting void was the worldwide pattern of availability. As indicated not substantially filled by anyone else until the mid- in table 4, Peru provided us with 52%-90% of our 1980s. In recent years, Chile has emerged as our major foreign supplier. We also import a modest but

4 fairly steady amount of meal (averaging about 1 30,000 MT annually) from Canada, and smaller and more variable amounts from miscellaneous other countries. Up until 1970, U. S. fish meal exports were negli- gible and tended to be disregarded in published sta- tistics. Exports have fluctuated widely from 4,300 to 77,400 MT during 1970-88. Exports exceeded imports in 1978, 1980, and 1983 (table 4). Total fish meal supply (;.e., production plus im- ports minus exports) has declined somewhat in the post-1972 period relative to earlier years (figure 2). The variability in supply closely parallels the vari- o!,,,,,,,,,,,,,,,,,,,,,,,,I,J ability in net imports (i.e., imports minus exports). 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 Domestic production (depicted by the difference be- YEAR tween supply and net imports) has been much more Figure 1. Fish meal exports by Peru, Chile, and all countries, 1960-87. stable by comparison.

126 THOMSON: US. MARKET FOR FISH MEAL AND OIL CalCOFl Rep., Vol. 31,1990

TABLE 3 800

Fish Meal Production in the United States by Species 700 Net imports (Thousands of Metric Tons) cn Year Menhaden Tuna-Mack Anchovy Other Total P 6oo 0 500 1960 198.1 24.0 0.0 41.0 263.2 K 1961 224.6 19.2 0.0 38.6 282.4 I- 1962 5 400 217.5 24.1 0.0 41.7 283.3 U 1963 167.1 24.5 0.0 40.6 232.2 E 300 1964 145.4 19.1 0.0 48.9 213.5 0 2 1965 159.7 23.0 0.0 47.8 230.5 u 200 cn 1966 122.5 23.0 4.1 53.9 203.4 3 1967 108.0 23.1 5.1 55.3 191.6 p 100 1968 129.9 26.1 2.5 54.7 213.3 c 1969 144.7 24.4 10.3 49.8 229.2 0 1970 171.1 24.2 14.7 34.2 244.2 1971 200.5 26.6 7.0 31.6 265.6 -lOOI,,,,,,,,,,,,,,,,,,,,,,,,,,, 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 1972 175.6 39.2 10.1 34.1 259.0 1973 171.3 39.6 20.0 22.4 253.2 YEAR 1974 185.0 43.7 12.8 23.1 264.6 1975 173.6 33.7 25.1 20.9 253.4 Figure 2. US. supply and net imports of fish meal, 1960-88. 1976 192.9 36.4 20.1 21.9 271.3 1977 175.4 36.1 17.3 19.6 248.4 1978 250.8 45.9 1.9 22.2 320.9 PRICES 1979 254.7 43.0 9.0 --.77 j 329.2 1980 246.0 42.6 7.1 26.6 322.3 High-protein oil meals, like fish meal, are traded 1981 209.4 42.8 9.3 19.9 281.3 in a very competitive international market. The soy- 1982 273.9 32.1 7.3 17.1 330.4 bean meal price is generally considered to be a lead- 1983 286.6 37.8 0.5 14.2 339.0 1984 285.7 33.7 0.0 15.3 334.7 ing indicator for fish meal and other oil meal prices. 1985 279.0 31.3 0.0 9.3 319.6 Figure 3 is a graph of U.S. soybean meal and men- 1986 268.8 33.7 0.0 5.6 308.1 haden meal prices, both corrected for inflation to 1987 303.4 38.3 0.0 8.0 349.6 1988 228.9 34.5 0.0 20.1 283.5 1988 dollars. The prices are highly correlated, with .~ the price differential largely attributable to the dif- References: U.S. Department of the Interior 1960-1970; U.S. Department of Comnierce 1971-1988. ference in protein content.

TABLE 4 U.S. Fish Meal Imports, Exports, and Net Imports (Thousands of Metric Tons) Imports by country of origin Exports Net - _.~ Year Peru Chile Canada Other Total imports 1960 61.7 19.1 28.1 10.8 119.7 - 119.7 1961 137.3 11.0 34.7 14.6 197.6 - 197.6 1962 168.9 8.3 38.8 12.9 228.9 - 228.9 1963 258.9 21.4 46.3 14.8 341.4 - 341.4 1964 315.7 11.7 49.7 21.2 398.3 - 398.3 1965 190.3 5.2 39.7 10.4 245.6 - 245.6 1966 250.9 81.1 39.6 34.6 406.2 - 406.2 1967 401.4 37.1 42.3 110.2 591 .0 - 591.0 1968 182.6 18.3 13.0 19.2 233.1 - 233.1 1969 99.8 19.6 19.2 4.7 143.3 - 143.3 1970 73.9 6.4 22.4 2.4 105.1 4.3 100.8 1971 181.1 0.0 52.3 23.5 256.9 9.2 247.7 1972 319.5 0.0 25.0 11.1 355.6 9.4 346.2 1973 37.9 0.0 22.3 1.9 62.1 33.3 28.8 1974 26.7 0.0 27.5 7.8 62.0 50.3 11.7 1975 68.5 7.0 30.8 1.1 107.4 10.7 96.7 1976 72.0 0.0 30.8 24.6 127.4 30.0 97.4 1977 14.3 2.0 22.5 35.2 73.9 32.7 41.2 1978 6.0 0.0 29.7 4.1 39.8 46.0 -6.2 1979 25.6 7.5 24.7 23.5 81.3 14.2 67.1 1980 6.0 0.0 22.0 16.9 44.9 77.4 -32.5 1981 0.0 24.3 22.0 7.6 53.9 42.6 11.3 1982 4.7 42.8 22.4 6.6 76.5 16.2 60.3 1983 6.5 23.4 20.9 10.8 61.6 70.2 -8.6 1984 0.0 43.5 21.4 10.8 75.7 18.3 57.4 1985 0.0 131.6 23.0 77.0 231.6 31.4 200.2 1986 12.4 105.6 16.3 33.8 168.1 34.9 133.2 1987 27.9 94.1 29.2 27.1 178.6 46.9 131.7 1988 46.7 25.4 32.0 16.3 1'0.4 68.0' 59.4 ~ - "Error in published statistics corrected per Steve Kopliii, NMFS, Washington, D. C.,pers. comm. References: U. S. Department of the Interior 1960-1970; U. S. Department of Commerce 1971-1988. THOMSON: US. MARKET FOR FISH MEAL AND OIL CalCOFl Rep., Vol. 31,1990

1000 , I 40 I I

35

I

looI,,,,,,,,,,I,,,,,I,~,,~,,,,,l 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 613 60 70 EO 87 YEAR YEAR

Figure 3. U.S. menhaden meal price (60% protein, bulk, f.o.b., East Coast/ Figure 4. U.S. per capita consumption of chickens and turkeys in 1960, 1970, Gulf plants) and soybean meal price (50% protein, bulk, Decatur, Illinois) in 1980, and 1987. 1988 dollars, 1960-88.

A notable feature of the graph is the 1973 price The situation with regard to table birds is quite increase, which resulted from a serious worldwide different (figure 4). Poultry consumption, measured shortage of oil meals. Several factors contributed to in ready-to-cook weight, has more than doubled this shortage, including (1) major failures of oil meal from 15.5 kilograms (kg) per person in 1960 to 35.3 crops around the world, (2) increases in fuel-related kg in 1987. This change, compounded by the in- production costs due to the Arab oil embargo, and crease in population over this same period, has re- (3) the collapse of the Peruvian anchoveta fishery. sulted in a dramatic increase in poultry production. Table birds slaughtered under federal inspection in- DEMAND creased almost fourfold from 3.1 million MT (live weight) in 1960 to 12.2 million MT in 1987 (U.S. Demand for Poultry Products Department of Agriculture 1960-1988). In the years since World War 11, the average Amer- ican diet has shifted away from grain products in Accommodation of Poultry Industry to Limited favor of more animal protein. This increased de- Fish Meal Supplies mand for protein is partially reflected in the shift The large increase in poultry production has not from farm production of chickens to the factory- been accompanied by a commensurate increase in style mass production of commercial broilers that fish meal usage. The poultry industry has accom- we see today. Approximately 80% of the fish meal modated itself to limited fish meal supplies by sub- consumed in the United States is used as an ingre- stituting other ingredients in poultry feed mixes, dient in poultry feed (Vondruska 1980). The final and by making technological changes to promote demand for poultry products is an indicator of poul- rapid growth of chicks. try feed usage and the demand for fish meal. Over 70% of the cost of producing chickens and U.S. egg production increased steadily from 61 turkeys, excluding processing and marketing costs, billion eggs in 1960 to 70 billion eggs in 1967. In the consists of feed (Vondruska 1980). As a result, small twenty years since 1967, annual egg production has changes in feed prices can have a major effect on not exceeded the 1967 production level. This level- total costs. U.S. poultry feed mixers are very so- ing of production is due to two offsetting factors: phisticated in their use of linear programming tech- (1) a decline in per capita egg consumption, and niques to determine least-cost combinations of (2) an increase in population. Because of increased ingredients (Hansen 1981/1982). They are also very productivity per layer, the stock of laying hens has quick to change feed composition in response to declined slightly from 295 million hens in 1960 to changes in prices (Huppert 1980; Thomson 1984). 280 millions hens in 1987 (U.S. Department of Ag- The role of fish meal in these linear programming riculture 1960-1988; Rogers 1978). These trends models is best understood by examining its nutri- suggest that total feed usage by laying hens has not tional contribution to poultry feed. All fish meal changed significantly since 1960. contains lysine and methionine, which are essential

128 THOMSON: U.S. MARKET FOR FISH MEAL AND OIL CalCOFl Rep., Vol. 31,1990

for the development and rapid growth of chicks. TABLE 5 These amino acids are not found in grain meals, U.S. Aquaculture Production, 1980 and 1985, by Species except for soybean meal, which contains high levels Group (MetricTom) of lysine. Lysine and methionine are also available in Species group 1980 1985 ~~ -~ synthetic form. The synthetic versions can be used Catfish 34,855 123,344 to obtain a proper amino acid balance in feed mixes Salmon 3,455 38.320 that do not contain fish meal (Titus and Fritz 1971; Crawfish 10.849 29,545 Trout 21,836 23,000 Vondruska 1980; Hansen 1981/1982). Baitfish 10,000 11,276 Including fish meal in the diet of laying hens re- Oysters 10,755 10,215 duces mortality by retarding the accumulation of fat Other finfish - 6,364 Other shellfish 391 1,111 in their livers (Ralph Ernst, USDA/UC Coopera- Total 92.141 243.675 tive Extension, Oakland, Calif., pers. comm.). Fish ~~ ~ meal also produces a significant growth response in Reference Rhodes 1988 table birds. Nutritional requirements for table birds depend upon a bird’s stage of growth, so feed com- TABLE 6 position varies accordingly. For chickens and tur- World Aquaculture Production in 1975,1980, and 1985, by keys, the maximum inclusion rate for fish meal is Species Group (Thousands of Metric Tons) about 8%-9% for starter rations and 7% for grower Species group 1975 1980 1985 rations. Higher rates than this tend to give a “fishy” Fiiifishes 2628 8 3206 8 5697 2 flavor to the final product. Desirable minimum in- Crustaceans 29 7 75 0 281 6 clusion rates are 1%-2% for starter rations and Mollusks 1961 2 3299 7 2885 7 0%- ~~ ~- ~~ 1% for grower rations (Vondruska 1980). Subtotal 4619 7 6381 5 8864 5 Seaweeds 8, other NA NA 3565 1 Thus, one way that feed mixers have been able to ~ ~~~ ~~ ~~ Total NA NA 12429 6 satisfy the increased demand for poultry feed in spite ~~ ~ ~~ ~- -~ ~~ of having smaller amounts of fish meal has been by Reference Rhodes 1989 substituting other ingredients. For table birds, they have reduced fish meal from maximum to minimum recommended levels in starter rations and elimi- million MT in 1975 to 8.9 million MT in 1985 (table nated fish meal entirely from grower rations. They 6). Currently about 10% of the world’s fish meal have also largely eliminated fish meal from layer production is used to feed farm-raised finfish and rations (Ralph Ernst, USDA/UC Cooperative Ex- shellfish (FA0 1989). By one estimate (Rhodes tension, Oakland, Calif., pers. comm.). These 1988), world aquaculture production will reach 22 changes are consistent with Kolhonen’s (1974) pre- million MT by the year 2000 and account for about diction that, “In the long run fish meal will be used 25% of the world’s aquatic harvest. This and other as a unique small-quantity ingredient in high-qual- similar projections suggest a long-term increase in ity feeds rather than as a high-amount protein demand for fish meal in aquaculture. source. 9, Fish meal use has also been reduced by technolog- Alternative Usesfor Fish Oil ical improvements resulting in shorter time to mar- The National Fish Meal and Oil Association sub- ket for chickens and turkeys. In 1960 it took mitted a petition to the U.S. Food and Drug Ad- approximately nine weeks to bring a three-pound ministration in 1986 requesting approval to use broiler to market; today it takes six weeks (Ralph hydrogenated and refined fish oils in products for Ernst, USDA/UC Cooperative Extension, Oak- human consumption. The FDA recently granted land, Calif., pers. comm.). As a result, feed require- approval for the hydrogenated oil, paving the way ments (including fish meal requirements) per bird for its use in products such as shortenings and pas- have declined. tries. However, because of U. s. Department of Ag- riculture standards for margarine, fish oil still FUTURE TRENDS cannot be used in margarines. The portion of the petition pertaining to refined Aquaculture Demand for Fish Meal fish oil is still pending. Unlike hydrogenated oil, Aquaculture production in the United States al- refined oil contains omega-3 fatty acids, which have most tripled from 1980 to 1985 (table 5). World pro- been shown to provide a wide variety of health ben- duction has also increased dramatically, from 4.6 efits (Pique 1986). Adding refined fish oil to products

129 THOMSON: US. MARKET FOR FISH MEAL AND OIL CalCOFl Rep., Vol. 31,1990

such as salad dressings could enhance their nutri- 6, tional value and marketability. However, the long- term prospects for this are uncertain, since (1) FDA approval may or may not be forthcoming, and (2) the technology necessary to address the problem of ran- cidity in refined oil is not well developed (Paul Bauersfeld, NMFS, Charleston, S. C., pers. comm.).

Japanese and South American Sardine Fisheries Japan has historically been a major world pro- ducer of fish meal. From 1960 through 1971 Japan produced 9%-15% ofthe world’s fish meal; its share of production increased to 15%-21% from 1972 through 1987. Until the mid-1980s Japan was also a 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 net importer of meal (table 7). YEAR Japan derives most of its fish meal from its sardine Figure 5. Landings of Japanese and South American sardine, 1965-87. (Sardinops melanosticta) fishery, which has produced two periods of high yield in this century. Japan’s take most ofthe harvest, the Soviet Union and South sardine landings increased through the early 1900s Korea have also participated in the fishery since the to a peak of1.75 million MT in 1935, then gradually late 1970s. declined to 9,200 MT by 1965 (Lluch-Belda et al., in The South American sardine (Sadinops sagax) press). Since 1965, landings have again increased fishery has experienced similar rapid growth in the dramatically (figure 5). Although Japan continues to past two decades. Annual sardine landings by Chile and Peru have increased from negligible amounts in the mid-1960s to approximately 5 million MT TABLE 7 (figure 5). World Production and Japanese Production, Imports, and Exports of Meals and Solubles from Animals of Given the record harvests experienced in recent Aquatic Origin (Thousands of Metric Tons) years, the sardine fisheries ofJapan and South Amer- ica warrant close attention. A change in the status of World ~~ ~ ~ Japan- ~ these stocks could significantly affect the availability Year producnon Production Imports Exports ~~ and price of fish meal. 1960 2,076 0 312 7 19 4 6.3 1961 2,580 0 362 2 31 3 4.9 1962 2,900 0 390 0 38 5 18.1 New Products from Reduction Species 1963 2,902 0 328 4 84 3 3.6 Efforts are ongoing to develop marketable prod- 1964 3,666 0 353 1 102 3 6.2 ucts for direct human consumption from species tra- 1965 3,615 0 344 5 112 6 13.1 1966 4,170 0 433 5 95 6 15.8 ditionally used for reduction. For instance, a 1967 4,660 0 420 3 86 8 11.3 processor in Virginia has been exploring the eco- 1968 5,060 0 500 5 150 3 6.8 nomic feasibility of making a marketable surimi 1969 4,750 0 594 2 108 0 183.0 1970 5,450 0 671 0 94 7 24.5 from menhaden (Malcolm Hale, NMFS, Charles- 1971 5,400 0 692 0 21 7 37.7 ton, S.C., pers. comm.). The Fishermen’s Cooper- 1972 4,320 0 735 9 56 8 38.6 ative in San Pedro, California, is test-marketing 1973 4,020 0 791 0 87 1 17.8 1974 4,570 0 773 7 74 5 31.3 canned sardines for human consumption. Although 1975 4,510 0 839 6 70 6 49.3 the ultimate outcome of these enterprises is uncer- 1976 4,890 0 745 9 59 5 49.0 tain, the expectation is that alternative uses will be 1977 4,575 4 857 2 181 1 37.5 1978 4,916 1 890 2 84 9 64.3 found for reduction species over the long term. 1979 5,089 9 895 6 101 6 57.7 1980 4,971 9 879 9 141 0 43.3 Americanization of the Alaska Pollock Fishery 1981 5,056 2 898 9 84 1 73.7 1982 5,394 1 1,004 1 44 3 135.7 Significant increases in U. S. harvesting of and 1983 5,282 6 1,133 5 95 1 79.6 processing capacity for the Alaska pollock (Thevayua 1984 6,097 7 1,262 7 61 6 135.3 chalcoyvnnzr?ia) have resulted in a drastic curtailment 1985 6,275 2 1,166 6 80 3 157.4 1986 6,661 3 1,179 1 I61 5 167.2 of foreign landings and joint venture operations in 1987 6,394 6 1,112 7 187 3 216.6 recent years. As indicated in table 8, landings of Reference Food and Agriculture Organization 1960-1987 Alaska pollock by foreign vessels declined from an

130 THOMSON: U.S. MARKET FOR FISH MEAL AND OIL CalCOFl Rep., Vol. 31,1990

TABLE 8 trade, the United States could reverse its long-stand- Landings of Alaska Pollock (Metric Tons) ing status as a net importer of fish meal and become Joint a net exporter. Year Domestic venture Foreign Total ~ ~~ ~~ 1977 323 0 1,009,826 1,010,149 1978 1,765 0 1,074,077 1,075,842 1979 2,551 0 1,047,150 1,049,701 LITERATURE CITED 1980 1,409 11,800 1,119,126 1,132,335 FA0 (Food and Agriculture Organization of the United Nations). 1960- 1981 1,741 58,950 1,117,455 1,178,146 1987. Yearbook of fishery statistics: fishery commodities. 1982 1,479 128,886 1,051,949 1,182,314 . 1986. The production of fish meal and oil. FA0 Fisheries Tech- 1983 1,382 283,104 973,050 1,257,536 nical Paper 142, 63 pp. 1984 10,894 444,256 1,032,249 1,487,399 -. 1989. GLOBEFISH Highlights, no. 1/89:28. 1985 42,109 614,337 851,870 1,508,316 Glantz, M.H. 1979. Science, politics and economics of the Peruvian 1986 59,160 904,111 352,682 1,315,953 anchoveta fishery. Mar. Policy 3(1):201-210. 1987 250,407 1,057,315 3,596 1,311,318 Hansen, T. 1981/1982. A model of the world fish meal market. Fish. 1988 570,285 826,564 0 1,396,849 Res. 1271-288. ~~ ~~ ~~~ ~ Huppert, D. D. 1980. An analysis of the United States demand for fish Reference U S Department of Commerce 1977-1988 meal. Fish. Bull. 78(2):267-276. Kolhonen, J. 1974. Fish meal: international market situation and future. approximate annual average of one million MT dur- Mar. Fish. Rev. 36(3):36-40. Lasker, R., and P. Smith. 1977. Estimation of the effects of environmen- ing 1977-85 to zero in 1988. Joint venture landings, tal variation on the eggs and larvae of the northern anchovy. Calif. which peaked at one million MT in 1987, are also Coop. Oceanic Fish. Invest. Rep. 19:128-137. expected to decline to zero in 1990, and domestic Lluch-Belda, D., R. J. M. Crawford, T. Kawasaki, A. D. MacCall, R. H. Parrish, R. A. Schwartzlose, and P. E. Smith. In press. World- landings are expected to increase commensurately. wide fluctuations of sardine and anchovy stocks: the regime problem. With the Americanization of the fishery and re- S. Afr. J. Mar. Sci. cent increases in fish meal processing capacity in Pique, G. G. 1986. Omega-3: the fish oil factors. San Diego: Omega-3 Project, Inc., 163 pp. shoreside plants and aboard U. S. factory trawlers, Radovich, J. 1981. The collapse of the California sardine fishery: what pollock meal is expected to become an increasingly have we learned? In Resource management and environmental uncer- large component of U. S. fish meal production. Ad- tainty, M. H. Glantz, ed. New York: John Wiley & Sons, pp. 107- 136. ditional impetus may be provided by the North Pa- Rhodes, R. J. 1988. Status of world aquaculture: 1987. Aquaculture cific Fishery Management Council, which is magazine buyer’s guide, pp. 4-18. currently considering a change in regulations to . 1989. Status ofworld aquaculture: 1988. Aquaculture magazine buyer’s guide, pp. 6-20. require full use of the resource. Should such an Rogers, G. B. 1978. Costs, prices and productivity in the poultry and amendment be adopted, it could lead to similar re- egg industries. Poultry and egg situation. Washington, D.C., U.S. quirements for other Alaska groundfish species. Dept. ofAgriculture, pp. 26-29. Thomson, C. J. 1984. A model of fish meal supply and demand in the In 1988, the United States produced approxi- United States, 1966-1981. NMFS, SWFC Admin. Rep. LJ-84-04, mately 15,000 MT of pollock meal from the offal 27 pp. generated in the preparation of surimi and fillets/ Thomson, C., C. Cooney, and J. Morgan. 1989. Status ofthe California coastal pelagic fisheries in 1988. NMFS, SWFC Admin. Rep. LJ-89- blocks (Vondruska et al. 1989). Assuming a fish meal 14, 23 pp. yield of 10% from round weight (Steve Koplin, Titus, H. W., and J. C. Fritz. 1971. The scientific feeding of chickens. NMFS, Washington, D. C., pers. comm.) and an Fifth edition. Danville, Ill. The Interstate Printers &. Publishers Inc. 312 pp. average annual harvest of 1.2 million MT, U.S. pol- U. S. Department of Agriculture. 1960-1988. Agricultural statistics. lock meal production could reach 120,000 MT an- Washington, D.C. nually. This would be a significant addition to the U. S. Department of Commerce, National Oceanic and Atmospheric Administration, National Marine Fisheries Service. 1971-1988. Fish- 200,000 to 350,000 MT of fish meal that we cur- eries ofthe United States. Washington, D.C. rently produce each year. U.S. Department of the Interior, Fish and Wildlife Service, Bureau of Much of the pollock meal produced in recent Commercial Fisheries. 1960-1970. Fishery statistics of the United States. Washington, D.C. years has been exported to Taiwan’s eel farms (Jerry Vondruska, J. 1980. Postwar production, consumption, and prices of Babbitt, NMFS, Kodiak, Alaska, pers. comm.). Fu- fish meal. IJLResource management and environmental uncertainty. ture increases in pollock meal production may also M. H. Glantr, ed. New York: John Wiley & Sons, pp. 285-316. Vondruska, J., R. Kinoshita, and M. Milazzo. 1989. Situatmn and out- be exported abroad rather than absorbed into the look for surimi and surimi-based food. NOAA Technical Memoran- U.S. market. Depending on the magnitude of this dum NMFS-SEFC-233, 16 pp.

131 HUNTER ETAL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Yol. 31,1990

BATHYMETRIC PATTERNS IN SIZE, AGE, SEXUAL MATURITY, WATER CONTENT, AND CALORIC DENSITY OF DOVER SOLE, MICROSTOMUS PACIFICUS

JOHN R. HUNTER, JOHN L. BUTLER, CAROL KIMBRELL, AND ERIC A. LYNN National Marine Fisheries Service Southwest Fisheries Center f! 0.Box 271 La Jolla, California 92038

ABSTRACT mento marcado en el contenido de agua del cuerpo Ninety-eight percent of the spawning biomass of y consequentemente, por un decrecimiento en la Dover sole, Microstomus pacijicits, in central Califor- densidad calorica por gramo de peso h~medo.Por nia waters live in a region of the continental slope ejemplo, la deiisidad calorica de un pez que mide between 640 and 1006 m (350-550 fath.) character- 275 mm y vive a 200-400 m de profundidad decrece ized by low oxygen concentrations (0.27-0.36 ml/l) de 86 kcal por gramo de peso htimedo (83% de agua) and cold temperatures (5.9OC-3.2”C). Juvenile Dover a 60.3 kcal (90% de agua) para un pez que mide sole settle on the continental shelf and gradually 440 mm de longitud y vive a 900 m de profundidad. move down the slope over their lifetime, reaching Las hembras pueden vivir hasta proximadamente the oxygen minimum zone as they become sexually 10s 53 afios de edad y 10s machos hasta 10s 58. El mature. Fifty percent of Dover sole in central Cali- contenido de agua parece estar relacionado con la fornia reach sexual maturity when about 31 cm long edad y el largo del individuo y con la profundidad a and about seven years of age. The ontogenetic la cual vive. movement down the slope continues after sexual maturity and is accompanied by a marked increase INTRODUCTION in water content of the body and a consequent de- Dover sole, Micvostomus pactjicus, are found from crease in caloric density per gram wet weight. For the Aleutian Islands in the Bering Sea to Baja Cali- example, caloric density decreases from about fornia (Eschmeyer et al. 1983). The U. S. fishery for 86 kcal per gram wet weight (83% water) for a 275- Dover sole occurs from Point Conception, Califor- mm fish living at 200-400 m, to 60.3 kcal per grain nia, to the Canadian border. Dover sole inhabit wet weight (go0/, water) for a fish 440 mm long depths ranging from about 55 to 1300 m. Older and living at about 900 m. Female Dover sole may live larger fish usually occur in the deeper portion of the as long as 53 years, and males 58 years. Water con- depth range, and younger and smaller fish in the tent appears to be a function of age as well as length shallower depths. A seasonal inshore migration has and depth. been described: fish move into deep water in the fall before the spawning season and into shallow water RESUMEN in the summer (Hagerman 1952; Alverson 1960; El 98% de la biomasa del desove del lenguado de Pearcy et al. 1977). Most individuals apparently re- las aguas de California central, Microstornus paclficus, main in the same general locality throughout their vive en un Area del talud continental entre 10s 640 y lives. Although longshore movements of up to 360 10s 1006 m caracterizada por bajas concentraciones mi (579 km) in seven years have been recorded, 97% de oxigeno (0.27-0.36 mL/L) y bajas temperaturas of tagged individuals were recaptured within 50 km (5.9OC-3.2”C). Los juveniles se establecen en la pla- of where they were tagged (Westrheim and Morgan taforma continental y gradualmente, durante su 1963). desarrollo, se van moviendo hacia el talud y a lo Large Dover sole from deep water are often ‘tiel- largo de 61, llegando a la zona de minimo oxigeno lied” (have flesh with an unusually high water con- cuando alcanzan la madurez sexual. El 50% del len- tent). This ‘tjellied” consistency limits the market guado alcanza la madurez sexual cuando mide value of fillets from large Dover sole because their aproximadamente 31 cm de longitud, a 10s 7 aiios de desirability is reduced (Hendricksen et al. 1986). edad. El inoviniiento descendiente a lo largo del Owing to the ontogenetic migration into greater talud durante la ontogenesis continua pasada la depths, and the extensive depth range, the demo- madurez sexual, y es acompafiado por un incre- graphic and physiological characteristics of Dover sole change strikingly with depth, age, and length. ~~~ ~~~~~ ~~~ [Manuscript received April 23,1990.1 Thus neither the dynamics nor the ecology of Dover

132 HUNTER ETAL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

sole populations can be properly analyzed or under- randomly sampled, and 25 females were assessed for stood without a careful evaluation of the relation gonad maturation. Four to six fish of each sex were between depth and key physiological and popula- weighed and frozen for later removal of otoliths and tion variables. tissue. for analysis of water content. The bottom The objective of this paper is to describe the rela- temperature (reversing thermometer on Nansen tionships between depth, length, age, sexual matur- bottle) and oxygen content (Winkler titration) were ity, water content, caloric density, and biomass of measured for 17 Nansen cast stations ranging from Dover sole. We also provide data on the temperature 183 to 1,280 m (100-700 fath.; figure 1, left, trian- and oxygen content of the habitat in which fish of gles). The total trawl catch of Dover sole was different length and age are found. Our analysis does weighed during 1987 and 1988. In 1988, up to 100 not include data for the summer months, when fish Dover sole were randomly sampled and weighed may have a shallower distribution (Alverson 1960). by sex; the maturation state of all ovaries was determined.

METHODS Age Determination Ages were determined from otoliths removed Sea Collections from 341 females and 64 males captured during Research trawl collections were taken at depths 1985-86 and from 154 females and 97 males captured between 69 and 1394 m (38-762 fath.) off the central in 1987. Left otoliths were embedded in epoxy resin California coast between Point Conception and Half and cut with a diamond wafering blade in a thin Moon Bay, California, during 1985-88 by NOAA- cross section through the nucleus from dorsal to Southwest Fisheries Center (SWFC) personnel ventral (Chilton and Beamish 1982). Thin sections (table 1). Trawl collections were opportunistic be- were mounted on microscope slides with Eukitt fore 1987. In 1987, samples were taken at 183-m mounting medium, polished, and read with a com- (100-fath.) intervals along transect lines; in 1988 a pound microscope. A typical otolith section is random sampling design stratified by depth was shown in figure 2. Counting procedures followed used (figure 1). those of Chilton and Beamish (1982). Each otolith In all years fish were sexed and measured for total was read independently by three observers without length. Before January 1987, fish were randomly knowledge of the length of the fish. Otoliths whose sampled from each collection until 25 females had readings differed by more than 10% among the read- been identified and their ovaries preserved for later ers were reread using the same protocol until the assessment of maturity; the females and some males readings agreed to within 10%. Estimated age was were frozen for subsequent extraction of otoliths the average of the individual readings. Parameters of and determination of water content as described be- the von Bertalanffy equation relating length and age low. During January-February 1987, either all of the were estimated by the simplex method (O’Neill Dover sole in the trawl collection or 100 fish were 1971).

TABLE 1 Sources and Numbers of Dover Sole Used for Analyses Number of positive Number of Dover sole used in various analyses trawl collections ~~~ ~~~ ~~~

~~~ Depth Sampling ~-Percent Size at Sexual Oxygen and Dates VPea N Min Max water Calories Age depth maturity ~~ ~~ ~~ 1213-12/12/85 OP 11 94 704 108 7 37 195 104 1 /14-2/24/86 PS 2 330 600 115 - 180 - - 315-317186 OP 8 52 500 165 18 14 210 - - - 512-5/4/86 OP 3 175 537 60 I7 62 74 1111-21 15/87 LT 49 99 705 265 - 2.55 3225 - 17” 2123-4/9/88 SR 51 38 602 - - - 3800 - - -~ ~~ ~~~ Total specimens 713 27 548 6504 104 ~~~ ~~ ~ ~- ~~~~~ ‘OP = opportunistic trawl samples, PS = port samples, LT = line-transect trawl samples, SR = stratified random trawl samples “Number of Nansen cast stations.

133 HUNTER ET AL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31; 1990

180N

IFN

16ON

35ON

U 123OW 122w 121ow 1 'W

Figure 1. Left, trawl stations occupied on the January-February 1987 groundfish cruise of the R/V David Starr Jordan. Right, trawl stations occupied on the February-March 1988 groundfish cruise of the R/V David StarrJordan.

Figure 2. Thin section of the left otolith of a 24-year-old Dover sole. Numbers indi- cate years.

134 HUNTER ET AL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

Biomass 96 We used the mean biomass in the area swept by trawls, and total area within three depth strata to 94 estimate Dover sole biomass. The survey area ex- tended from 34"30'N, 120"30'W to 36"30'N, 92 122"30'W (Butler et al. 1989). -X I u, 90 Sexual Maturity Y The size at first maturity of females taken in De- cember 1985 was estimated by logistic regression analysis (Draper and Smith 1981; Engelman 1988). K k 66 Data from the other surveys (table 1) were not ap- s propriate for estimating sexual maturity because the 8 spawning season began before the first survey. The 84 possibility of mistaking immature ovaries for post- spawning ovaries was minimized because Decem- 82 ber appears to be early in the spawning season (Hunter, unpublished data). Ovaries with yolked 80 oocytes were considered to be mature, and those I without yolked oocytes to be immature. 80 82 84 86 88 90 92 94 96 % WATER WHITE MUSCLE (h,) Water Content We measured water content by determining the Figure 3. Relation of water content of whole Dover sole to water content of white muscle. Solid line is regression line fit to data; dashed line is one-to- wet weight of a tissue sample (about 3.5 g) and then one relationship. drying it to constant weight at 60°C. The white muscle samples were taken from the right side of the fish between the lateral line and the dorsal fin at the insertion of dorsal rays 30 to 36, and the red muscle Water content of the red muscle tissue sample was samples were taken from the right side above the also linearly related to water content of the entire lateral line and behind the eye. fish, although the relationship was poorer than that To determine whether a single tissue sample rep- for white muscle: resented water content of the whole fish, we mea- sured the entire water content of 20 females, H = -51.1 + 1.62 h2, (2) 6 males, and one fish of indeterminate sex by grind- ing and drying the entire fish after removing red and where h, is water content of red muscle (v2 = 0.837, white muscle tissue samples from the locations de- n = 27; p < 0.01). scribed above. We then regressed water content (in percent ofwet weight) ofthe entire fish (H)on tissue Caloric Density and Fat Content sample water content (figure 3): To determine caloric density (kcal/gm) of entire fish for 26 Dover sole, we used a Parr bomb calori- H = -5.08 + 1.05 h,, (1) meter and standard techniques (Parr Instrument Co. 1960; Paine 1971). The values reported here are the where h, is water content of white muscle (r2 = means of three determinations. Fish were selected 0.906, n = 27; p < 0.01). The slope of this equation by size to obtain caloric measurements over a wide did not significantly differ from 1.0, nor did the in- range of water content (the fish sampled ranged tercept differ from 0. On the basis of these results from 80% to 94% water). we assumed that the percentage of water in the white To determine fat content we used a Soxhlet ex- muscle tissue samples was the same as that of the traction with 2:l chloroform-methanol. This tech- whole fish. Whole fish water content was estimated nique did not provide an accurate estimate of total (equation 1) from white muscle tissue samples col- fat content because the extraction period (48-72 hrs) lected at sea for 106 fish collected in 1985, 335 fish was too short for fish with a high fat content, such collected in 1986 (go0/, females), and 265 fish (males as Dover sole. Notwithstanding this bias and other and females) collected in 1987. problems associated with chloroform-methanol ex-

135 HUNTER ETAL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

tractions (Dobush et al. 1985), we believe that the data provided useful information about fat concen- tration. 500600 1 - RESULTS E 400 E Y E 300 Length and Depth (7 Z The mean length of Dover sole increases rapidly W J 200 with depth over the first 300 fath. (549 m) and there- after more slowly. The relationship between depth and size of Dover sole varied little among years (figure 4, top and middle). The combined data for loo 1 04 all surveys clearly indicate that males taken at a given 0 20 40 depth are smaller than females (figure 4, bottom). --- I I 500 - Age and Growth Male and female Dover sole differ in size at age I E 400- E (figure 5). Estimates from counts of annuli in thin v sections of otoliths indicate that the fish live long 300- lives. The maximum estimated age was 56 years for (7 a 460-mm male and 51 years for a 492-mm female. FEMALE DOVER SOLE

o! I E 0 20 40 60 400 /+-- F AGE (years) 9 300 Figure 5. Top, length and age (calculated from otolith section) of male Dover Y sole. Solid line is von Bertalanffy curve with L- = 437 mm, growth parameter z K = 0,089, and to = ~ 4.7yr. Bottom, length and age (calculated from otolith 3 200 1988 a_--- I c section) of female Dover sole. Solid line is von Bertalanffy curve with L% = L 1 474 mm, growth parameter K = 0.085, and to = - 5.5 yr. See text for equa- 100 tion 3. 0 200 400 600 800 500 I

k 400 Parameters of the von Bertalanffy growth model, E A- a 300 DOVER SOLE MALES !I LENGTH BY DEPTH z 1985-86 - 200 1987 c - - I 1988 e---- I were L, = 437 mm, K = 0.089, and to = -4.7 100 J I 0 200 400 600 800 years (n = 161, p < 0.01) for male Dover sole. Esti- 500 1 i mates ofparameters for females were L, = 474 mm, -E 400 - K = 0.085, and t,, = -5.5 years (n = 495, p < F 0.01). Male and female Dover sole longer than 9 300 - 40.0 cm may be any age from 8 years to 40 or 1 50 years. Neither our methodology nor any other $ 200 - has been validated for Dover sole. Nevertheless, we 2 believe our age estimates approximate the true age, 100 -I I 0 200 400 600 800 because the otolith section method has been vali- DEPTH (fathoms) dated for another species in the same habitat (Ano- 0 200 400 600 800 1000 1200 1400 ploponiajirnbvia; Chilton and Beaniish 1982). DEPTH (meters) Our age estimates are much greater than those

Figure 4 Mean length of Dover sole collected at different depths Top, fe- obtained from scales (Demory 1972; Mearns and males, middle, males, bottom, all years combined with ?2 SE Harris 1975). However, tag-and-recapture studies

136 HUNTER ETAL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

(Pikitch and Demory 1988) have shown that scales figure 7). The water content of Dover sole (both underestimate the age of Dover sole. The maximum male and female) as a function of length showed age observed in this study (56 years) is in agreement little variation between years; the data for December with the longevity that Pikitch and Demory (1988) 1985-May 1986 were essentially the same as those predicted from tag-and-recapture studies and errors for January-February 1987 (figure 7). Combining in scale readings. A significant fraction (40%) of data for all years and sexes indicates that the average females in our samples were older than 20 years. water content for fish less than 275 mm was constant Our results suggest that estimates of the productiv- at 82.9% (n = 90, SD = 1.00), and for fish greater ity of Dover sole stocks based on ages estimated than 275 mm it increased with length (L)according from scales may be seriously flawed. to the equation

Water Colztent H = 72.7 + 0.036 L (4) Data for females taken from December 1985 through May 1986 indicate that the mean water con- (v2 = 0.58, n = 506; p < 0.01). tent of white muscle remains constant at about 82% in 15-30-cm females but increases with length Water Content, Age, Length, and Depth in larger Dover sole to about 90% in 50-cm females In the previous sections we showed that water (figure 6). The concentration of water in the red content increases with fish length and that length muscle followed a similar trend, but the increase in increases with depth and age. In this section we de- water content with length was much less. Average scribe the relationships between water content, water content of red muscle in 50-cm females was length, depth, and age of Dover sole females. A only 85%. These data indicate that red muscle is conserved. The red muscle in the anterior upper trunk region of the eyed side of the body is the larg- est concentration of red muscle in the body and is probably used to control head movements during 1987 feeding. The color of the red muscle changed with fish length. Larger fish had darker red muscle, indi- cating that the myoglobin content of the muscle Male,? may be higher in larger fish. Female The mean water content of males taken in Janu- ary-February 1987 did not differ from that of fe- males from that sample within the same 5-cm length class (ANOVA for five 5-cm length classes, 254-

324 to 475-524 mm, per sex where p = 0.90; I I I I 80 '

ni n W 92 - I- -I90 DOVER SOLE - -1987 - 89 90 - *-- 1985-1986 E 88. - I- 88 - 1985-1987 Z 87. w - c z 86- 0 86 o 85. a w 84. 84 I- $ 83' 82 82 . w 80 500 600 81 t 1 zz 100 200 300 400 80 1 I I I I I I I I 100 150 200 250 300 350 400 450 500 TOTAL LENGTH (mm) MEAN TOTAL LENGTH (mm) Figure 7. Top, water content of white muscle versus total length of female Figure 6. Water content in white and red muscle versus total length of female (N = 163) and male (N = 98) collected in 1987. Bottom, water content in Dover sole. Insert shows locations of tissue samples-W for white muscle white muscle of both sexes for 1985-86 (dashed line), 1987 (thin line), and and R for red muscle. 1985-87 (thick line).

137 HUNTER ET AL.: BATHYMETRIC PATTERNS OF DOVER SOLE ColCOFl Rep., Vol. 31,1990

stepwise multiple regression analysis indicated that on physiological grounds because of our failure to all three independent variables (age, length, and extract all the lipids in some of the fish. Neverthe- depth) accounted for significant variation in the de- less, figure 8 clearly indicates that caloric density and pendent variable, water content: the coefficient for fat concentration are linked, as would be expected. age was the first selected (table 2). The final No significant relationship existed between ca- equation was loric density and water content. There was no ap- parent relationship between fat concentration and H = 78.7 + 0.1226 T + 0.0123 L + 0.0026 D (5) water content (v’ = 0.02) when length was not in- cluded as a variable. When length (L)was included where H is water content (percent of wet weight), as an additional third dependent variable in a step- T is age (years), L is length (mm), and D is depth wise multiple regression analysis (table 3), a definite (fath.) (R2 = 0.69, n = 519; p < 0.01). Biological relationship existed between caloric density (C,) and interpretation of the significance of the individual regression coefficients in this equation could be mis- leading (Sokal and Rohlf 1981) because, as is indi- cated in the correlation matrix (table 2), all the + 7’5 I I variables are correlated with each other; length is P 7- correlated with age and depth as well as water $ a content. > a a 6.5 - ma W Caloric W a a. Density U Caloric density (kcal per g ash-free dry weight) $6- a a. 0 :.a was correlated with fat concentration (F,in percent a a a of dry weight). This relationship was less precise u a - a (Y’ = 0.57, n = 26; p < 0.01) than one would expect 2 5.5 s a

TABLE 2 10 20 30 40 50 60 Analysis of the Relation Between Water Content (H)of FAT CONCENTRATION (“A) Dover Sole and Their Age (T),Length (L),and Depth Figure 8. Caloric density versus fat concentration in Dover sole. (D) Stepwise regression Step 1 2 3 Constant 82.63 78.31 78.71 TABLE 3 Age (T) 0.2213 0.1272 0.1226 Analysis of Variance Stepwise Regression of the Caloric t-ratio’ 28.28 10.94 10.69 Length (L) 0.0157 0.0123 Density of Dover Sole (C,, kcal/g ash-free dry weight) t-ratio 10.25 7.30 on Their Total Length (I., in mm) and Water Content Depth (D) 0.00260 (H,in Percent) 4.61 t-ratio Summary S 1.82 1.66 1.63 Step 1 2 RZ 60.73 67.37 68.62 Constant 8.755 13.540 Analysis of variance Water (H) -0.032 -0.104 Source DF ss MS F P t-ratio’ -1.65 - 5.86 Regression 3 2,981.00 993.67 375.31 <0.0005 Length (L) 0.0042 Error 515 1,363.52 2.65 t-ratio 5.76 Total 518 4,344.52 S 402 263 R2 10.21 63.22 Source DF SeqSS Analysis of variance Age (T) 1 2,638.47 Length (L) 1 288.64 __Source - DF ss MS F P Depth (D) 1 53.89 Regression -7 2 725 1363 19 76

138 HUNTER ET AL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

water content of white muscle (H,in percent wet We also wished to examine how calorie content weight) yielding the equation varied with depth. We used the multiple regression equation 6 to compute the caloric density (C, in kcal C, = 13.540 + 0.0042 L - 0.1044 H (6) per g ash-free dry weight) of female Dover sole as a function of length and water content. We grouped with R2 = 0.60, n = 26; p < 0.01 (figure 9). To the estimated caloric densities into seven 100-fath. further evaluate these data we analyzed covariance (183-m) depth classes and calculated a mean and by arranging the data into three length classes (114- standard deviation (SD) for each class; the SD in- 296 mm, 303-396 mm, and 408-506 mm) and re- cludes the variance associated with equation 6 gressed water content on calories within length (Draper and Smith 1981; figure 10, top). class. This analysis indicated that the relationship A significant difference existed between the mean between calories and water content differed signifi- caloric density of females in the 100-fath. depth class cantly among the three length classes (233, 350, and and those in all other depth classes combined (t = 453 mm mean total length; p < 0.01). For the overall -7.34, d.f = 37.7, p < 0.001; d.f was computed mean water content (85.6'/0), the adjusted mean kcal from the formula given by Zar [1984]). The mean per g ash-free dry weight are 5.634 at 233 mm, 6.078 caloric density for females taken in the 100-fath. at 350 mm, and 6.424 at 453 mm. The relationship depth class was 5.761 kcal per g ash-free dry weight between caloric density and length was not signifi- (n = 34, SD = 292) and that of females taken at cant (Y' = 0.037, y1 = 27; p = 0.169). Thus, the relationship between caloric density and water con- tent changes with length. The relationship between length, water content, and caloric density in terms of wet weight (C,) was also evaluated. The multiple regression equation was

C,, = 651 + 0.0742 L - 7.06 H (7) where C, = kcal per 100 g wet weight, L = length, and H = water content of white muscle (R = 0.98, n = 26; y < 0.001). R2 is high because the water content of the fish largely determines the caloric ".T density when it is expressed on the basis of wet 0 100 200 300 400 500 600 700 DEPTH (fathoms) weight. DEPTH (meters) 200 400 600 800 1000 1200 0.10 20 I ." - C = 13,540- 104H + 4.2L 1 L = 114 - 296mm 6.8 - 0.09 18 0 0 L = 303 - 396mm I CT + L = 408 - 506mm W I 6.6 - P I- wP g 0.08 16 $ 6.4 3 - I- a:t L z n 6.2 - 3 0.07 14 UI W w a 2 6.0 - 0 w : ? 0.06 12 4 2 5.8 - 0 I 20 z (3 5.6 - 2 0 0.05 10 Y 5.4 - I 0.04 8 5.2,,,,,,,,,,,,,,,, 0 100 200 300 400 500 600 1 DEPTH (fathoms) PERCENT WATER (H) Figure IO. Top, kcal perg ash-free dry weight, SD and Nwithin 100-fath. depth Figure 9. Caloric density and water content of Dover sole in three size classes. classes. Arrow is mean caloric density for depths >150 fath. Bottom. kcal per (See equation 6 in text for multiple regression relation of caloric density with 100 g wet weight (left axis) and the complement of water content (right axis) length and water content.) within 100-fath. depth classes.

139 HUNTER ET AL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol:31,1990

depths greater than 150 fath. was 6.141 kcal (n = - 426, SD = 272). Thus, the Dover sole living in 1.0 shallow water (50-150 fath.; 91-274 m) have a lower caloric density per unit dry weight than Dover sole 0.9 - //- living at greater depths. 'a / / - A different pattern in the change in caloric density 0.8 .lo with depth occurs if caloric density is expressed in v) terms of wet weight rather than dry. Using the same 0.7 - a HAGERMAN 1952 data and equation 7, we recalculated the average ca- E - loric density of females on a wet-weight basis per U. 0.6 W depth class. On a wet-weight basis, the caloric den- a 3 sity (kcal per g wet weight) declines with depth be- + 0.5 - cause of the increase in water content (figure 10, 2 ----if-20. I - 7( I - I bottom). As would be expected, the decrease with 0 0.4 !- I depth in caloric density of Dover sole wet weight 0 I mimics the decline in the complement of the water 2 0.3 - I LL I content (100 - percent water; figure 10). I These measurements were made during the pe- 0.2 - I riod in which Dover sole mature sexually and begin / I I I to spawn (December-February). Other patterns 0.1 ~ I may exist at other times of the year. 11 I I I I 0.0 t 1 I I I I 1 200 250 300 350 400 450 500 i311 363 Sexual Maturity MEAN TOTAL LENGTH (mm) The sexual maturity of Dover sole females was estimated by calculating the fraction mature (n/r) for Figure 11. Circles, observed fraction of Dover sole that are mature in Decem- ber within 50-mrn length classes with N indicated, solid line, calculated frac- each of seven 50-mm length classes (255-526 mm). tion mature from logistic regression equation (equation 8); Hagerman line These data were fit by maximum likelihood esti- calculated similarly; dashedline, length at 50% maturity. mates of the parameters to a logistic model

Oxygen Minimum Zone Perhaps the most striking features of the environ- where a = -9.947, standard error (SE) of a = ment occupied by Dover sole are the oxygen mini- 0.00688, and t = 4.657 (d.f = 102, p < 0.01); mum and dysaerobic zones. In the 1987 survey the b = 0.0320, SE of b = 2.159, and t = -4.608 (d.f. oxygen minimum zone (0, < 0.5 ml/l) occurred = 102, p < 0.01); and n = 104. This equation pre- between 640 m (350 fath.) and 1010 m (550 fath.), dicts that 50% of Dover sole females of 311-mm which is similar to the depth range (280-550 fath.) length are mature (figure 11). The smallest Dover during 1981-82 reported by Mullins et al. (1985) off sole with advanced yolked oocytes in our collections Point Sur, California. The dysaerobic zone (0,< 1 was 290 mm long. Judging from our age-length re- ml/l) began at 457 m (250 fath.). Most sexually ma- lation for female Dover sole (equation 3), a length of ture females were observed in the dysaerobic zone. 31 1 mm corresponds to an age of about 7 years, and We estimated that 86% of the spawning biomass a length of 290 mm corresponds to about 6 years. existed in the oxygen minimum zone (table 4; figure The mean lengths per depth class were used to 12) on the basis of the middle depth stratum (250- calculate how sexual maturity varied with depth. 549 fath.) used in the 1987 and 1988 surveys (Butler Mean female lengths were computed for each of et al. 1989). We also estimated that nearly all (93%) eight 100-fath. (183-m) depth classes (all trawl data, of Dover sole living above the dysaerobic zone were 1985-88; n = 4,412; table 1; figure 4), and the ma- juvenile. turity was calculated using equation 8. This analysis The oxygen minimum zone is also characterized indicated that sexual maturity increased with depth by low temperatures (figure 12). The water in the from about 60% at 200 fath. to 90% or more at dysaerobic zone ranged from 5.9"C at 530 m (290 depths of300 fath. (549 m) or greater. fath.) to 3.2"C at 1262 m (690 fath.), and that in the

140 HUNTER ETAL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

DEPTH (Fathoms) 0 200 400 600 800 9

7 -u g5 3 00 u3 ml TEMPERATUR 3 2 3 2 m OXYGEN

a r. 00 2 m-

00 0 r. PIr. r.4 am b. 0 .% p 'f I mu3 0 2 5

w

L u 12 0 3 r. 8 10 dW r. 7 2 2 1 a 05 voY 200 400 600 800 0d. DEPTH (Fathoms) 0" 3

IIII...1111111, 0 200 400 600 800 1000 1200 1400

DEPTH (Meters) icPI

Figure 12. Bottom temperature, dissolved oxygen, biomass (MT) in study a area, length (mm), and energy content (kcaV100 g wet weight) of Dover sole as a function of depth for 1987.

PI ic m PI 3 d ic 3 2 oxygen minimum zone from 4.9"C (700 m; 383 PI fath.) to 4.3"C (902 m; 493 fath.). Reduced metabo- f icm d W d lism due to temperature may help Dover sole cope 2 K with the low oxygen concentrations in the region.

In 0 ic r. PI Wic DISCUSSION PI d c Mu tu vity Our estimate ofthe length at first maturity differs from that estimated by Hagerman (1952). We fit equation 8 to Hagerman's data (250-630 mm, tz = 846) and estimated the length at 50% maturity of female Dover sole landed in Eureka, California, during 1948-49 to be 363 mm (u = - 17.85, SE of u = 1.975, t = -9.042, d.f = 842, p < 0.01; b = 0.0492, SE of b = 0.0050, t = 9.829, d.6 = 842, p < 0.01; and n = 844). We compared the coeffi- cients of the logistic regressions for our data to those collected by Hagerman using the Z-test (Zar 1984;

141 HUNTER ET AL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

TABLE 5 Comparison of the Coefficients for Logistic Length and Maturity Equations for Female Dover Sole a Variance Z b Variance Z Present study -9.947 4.661 2.702 0.03202 0.000047 -2.025 Haeerman (1952) - 17.854 3.901 0.04921 0.000025 table 5). Because both sample sizes were large, we tured at a smaller size than those in central Califor- used the normal deviate Z with critical value of nia. For example, all Oregon females longer than > 12 I. The regression coefficients are significantly 320 mm were mature, whereas in central California, different at the .05 level, assuming that these distri- only 57% (SD = 8%) of 320-mm females were butions are normal. Thus it appears that Dover sole mature. Yoklavich and Pikitch (1989) suggest that females from central California matured in 1985 at a the long-term effects of the size-selective trawl fish- smaller size than did those from Eureka, California, ery off Oregon was a compensatory decrease in the in the 1940s. For example, 50% of female Dover sole size of first maturity of Dover sole. This hypothesis from central California mature when they reach 311 may also explain the difference between our esti- mm, whereas 50% maturity occurred at 363 mm in mates for central California and theirs for Oregon, females from Eureka (1948-49). This 52-mm dif- since the central California fishery (Morro Bay) ference may be equivalent to an average difference for Dover sole is a new fishery with only minor land- in 'the age of first maturity of four to five years ings until the last four or five years, whereas the (assuming fish from central California in the 1980s Dover sole fishery in Oregon has been active for grew at about the same rate as fish from Eureka in over forty years. the late 1940s). Similarly, Yoklavich and Pikitch (1989) compared their 1985 maturity estimate for Dover sole from High Water Content Oregon to one made for the same area 35 years ear- High water content (jellied flesh) has been re- lier. They found all females greater than 320 mm ported for four marine flatfishes: Dover sole (Fisher were mature in 1985, whereas 35 years earlier (Harry et al. 1987; Puckett 1989); winter flounder, Pseudo- 1959) only 50% of Oregon females of 380 mm were pleuronectes amevicanus (Pearcy 1961); yellowfin sole, mature and only 15% of the Dover sole less than 380 Limundu uspevu (Kizevetter et al. 1965); and Ameri- mm were mature. Yoklavich and Pikitch concluded can plaice, Hippoglossoides platessoides (Templeman that Dover sole from Oregon presently mature at and Andrews 1956). Roff (1982, 1983) attributed smaller size and probably younger age than they did high water content in flatfishes to degradation of 35 years ago. muscle tissue during periods of gonad development Because of possible sampling biases, we hesitate and maintenance during winter fasting (Roff 1982, to attribute the differences between recent and older 1983). On the other hand, Puckett (1989) concluded estimates to real biological differences. Bias could after examining the water content of Dover sole that result from failure to obtain a representative sample depth, annual reproductive cycles, size, age, and the over the full bathymetric range of Dover sole. In onset of sexual maturity were all involved, but addition, the two older estimates (Hagerman 1952 depth was the most important factor. Most signifi- and Harry 1959) may be biased because samples cantly, Puckett's analysis indicated that reproductive were taken from the fishery during spawning sea- condition of mature females living at 650-1020 m son. This process could lead to an overestimate of was not correlated with their water content. He the average size of fish at first maturity, because fe- found that the average water content of males and males in postspawning condition could have been females from deep water remained high throughout wrongly classified as immature. On the other hand, the year; for example, quarterly means for females the methods employed by ourselves and Yoklavich ranged from 89.6% in the winter to 91.6% in the and Pikitch (1989) were similar; in both studies the spring. He concluded that the effect of the reproduc- samples were taken in December 1985, which is tive season on water content was minimal, and that early enough in the spawning season that biases movement into a deepwater habitat was the key from misclassification of postspawning fish seem variable. unlikely. If our samples and those of Yoklavich and Dover sole differ from other flatfishes with high Pikitch (1989) accurately represent their respective water content in two ways: the adult population regional populations, then females in Oregon ma- lives at considerable depths (800-1500 m) and in the

142 HUNTER ET AL.: BATHYMETRIC PATTERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

oxygen minimum zone. In deep-sea fishes, the ca- concerning the ontogenetic movements we have de- loric density per g wet weight typically decreases scribed. Do the fish precisely re-sort themselves by with depth (Somero et al. 1983). The change in ca- depth each fall as they leave their shallow summer loric density of Dover sole with depth follows a habitat? How do they gradually (over decades) similar pattern (figure 12). The water content of increase the depth of their winter habitat? Does deep-living Dover sole is at the high end of the range high water content affect the extent of seasonal for the deep-sea fishes examined by Childress and movements? Nygaard (1973), where 90.5% was the highest value No data exist to fully answer these interesting recorded. This indicates that the ontogenetic change questions, but some inferences can be made on the in water content in Dover sole may be simply an basis of a tagging study conducted by Quirollo and adaptation to deepwater existence, and the explana- Kalvass (1987). Their data indicate that all Dover sole tions proposed for low caloric density of deep-sea may not participate in the summer inshore move- fishes may apply equally well to Dover sole. These ment. Mature fish tagged and released in shallow include the scarcity of food and development of water were usually recovered from deep water in the feeding strategies that permit a great reduction in winter and fall and from shallow water in the spring propulsive systems demanding a high metabolic rate and summer, indicating annual inshore movement. (Somero et al. 1983). In contrast, most of the mature fish tagged and re- Another plausible explanation for high water con- leased in deep water were recovered in deep water tent is one linked to life in the oxygen minimum regardless of season. Thus the tagging data indicate zone. The oxygen and energy required to maintain that two substocks may exist, one that migrates and white muscle in the oxygen minimum zone could one that does not. We suggest that the fish compos- be significant. Reduction of white muscle content ing the migratory substock may be younger and would reduce basal oxygen demand and increase have a lower water content than those composing the scope for activity in an oxygen-limited en- the nonmigratory substock. vironment. In contrast to white muscle, red muscle tissue is conserved in Dover sole. This red muscle is located CONCLUSIONS behind the head and at the base of the pectoral fin. The deep, cold, and poorly oxygenated region of This location may indicate a role in feeding. Feeding the continental slope known as the oxygen mini- behavior has been described for the congeneric mum zone is the habitat for the mature Dover sole lemon sole, Microstomiis kitt, by Steven (1930). This in central California. Ninety-eight percent of the species feeds on polychaetes and also inhabits spawning biomass of Dover sole occurs in this re- muddy bottoms. When feeding, a lemon sole raises gion. Dover sole spawn at these depths, and their its head and tail and sits perched on its side, scanning eggs rise to the surface layers. the substrate. When a polychaete is found, the fish Juveniles settle on the continental shelf and, with pounces with a forward leap, bringing its head down sexual maturity, gradually move down the conti- on the prey and strongly arching the anterior body nental slope. The onset of sexual maturity and the (Steven 1930). Dover sole also feed on polychaetes ontogenetic movement into the cold oxygen mini- (Pearcy and Hancock 1978; Gabriel and Pearcy 1981; mum zone (350-550 fath.; 640-1010 m) is usually and Wakefield 1984), and we have observed similar associated with an increase in water content, myo- feeding behavior in captive Dover sole in our labo- globin content of red muscle, and fat stores. The ratory. Dover sole perched on the substrate have also movement down the slope corresponds with a con- been observed by Allen (1982) and Wakefield (pers. sistent pattern of increasing size, age, and water comm., October 1988). Conserving red muscle content. while sacrificing white muscle as the fish becomes The ontogenetic movement down the shelf is more watery may be a mechanism to preserve levels gradual and occurs over decades. The average fe- of feeding performance while reducing basal oxy- male Dover sole in central California reaches matu- gen demand. rity when about 7 years old and 311 mm long. At this time she lives at a depth of about 329 m (180 Ontogenetic and Seasonal Movements fath.), and her water content is about 83.8%. We Tagging studies (Westrheim and Morgan 1963; speculate that over the next nine years the annual Quirollo and Kalvass 1987) and fishery data (Alver- inshore and offshore movement gradually ceases. son 1960) indicate that Dover sole move inshore in Over the same period the water content of the aver- the summer. This raises some interesting questions age female gradually increases as she descends to

143 HUNTER ETAL.: BATHYMETRIC PATERNS OF DOVER SOLE CalCOFl Rep., Vol. 31,1990

greater depths and enters the oxygen minimum zone Hendrickson, G. L., R. A. Fritzsche, and H. M. Puckett. 1986. Ecology (640 m; 350 fath.). By then she is 16 years old, is and possible causes of the ‘>ellied”condition in Dover sole, Microsto- mus pacificus. California Sea Grant biennial report, 1982-1984, almost 400 mm long, and has a water content of University of California Sea Grant College Program, Rep. No. 87%. By this time she, like 93% of her cohort, is R-CSGCP-020:129-130. sexually mature, and growth has slowed from 14 Kizevetter, I. E., E. F. Kleie, A. A. Kirillova, 0. M. Mel’Nikova, V. M. Myasoedova, and L. Ya. Ertel. 1965. Technological characteristics of mm per year when she was 7 years old to 6 mm per Bering Sea fishes, in Soviet fisheries investigations in the northeast year. Eleven years later the average female has de- Pacific. Part VI, €? A. Moiseev, ed. (Transl. from Russian by Israel scended to 1006 m (550 fath.), is 27 years old, has Program for Sci. Trans., Jerusalem, 1968.) Pp. 191-258. Mearns, A. J., and L. Harris. 1975. Age, length, and weight relation- grown 44 mm (about 4 mm per year), and has a ships in southern California populations of Dover sole. S. Cal. Coast. water content of 88.7%. Water content continues to Water Res. Proj. TM 219, 17 pp. increase slowly over the next decades, and the fish Mullins, H. T., K. McDougall, and T. L. Vercoutere. 1985. Oxygen- minimum zone edge effects: evidence from the central California continue to spawn and move deeper. The oldest fe- coastal upwelling system. Geology 13:491-494. male we aged was 51 years. The highest water con- O’Neill, R. 1971. Function minimization using a simplex procedure. tent was 94.1%. The greatest depth at which we Algorithm AS47. Applied Statistics, 338. Paine, R. T. 1971. The measurement and application of the calorie to collected Dover sole was 1269 m. ecological problems. Annu. Rev. Ecol. Syst. 2145-164. Parr Instrument Co. 1960. Oxygen bomb calorimetry and combustion methods. Technical Manual No. 130, 1-56. Moline, Ill. Pearcy, W. G. 1961. Seasonal changes in osmotic pressure of flounder LITERATURE CITED sera. Science 134:193-194. Allen, M. J. 1982. Functional structure of soft-bottom communities of Pearcy, W. G., and D. Hancock. 1978. Feeding habits of Dover sole, the Southern California Shelf. Ph.D. thesis, Univ. Cal. San Diego, Microstomits pacificus; rex sole, Glyptocephalus zachirus, slender sole, 577 pp. Lyopsetta exilis; and Pacific sanddab, Citliarichthys sordidus, in a region Alverson, D. L. 1960. A study of annual and seasonal bathymetric catch of diverse sediments and bathymetry off Oregon. Fish. Bull., U.S. patterns for commercially important groundfishes of the Pacific 76641-651. Northwest coast of North America. Pac. Mar. Fish. Comm. Bull. 4, Pearcy, W. G., M. J. Hosie, and S. L. Richardson. 1977. Distribution 66 PP. and duration of pelagic life of larvae of Dover sole, Microstornus paci- Butler, J. L., C. Kimbrell, W. C. Flerx, and R. D. Methot. 1989. The ficus; rex sole, Glyptocephalrrs zachivttr; and petrale sole, Eopsetra jor- 1987-88 demersal fish surveys off central California (34”30’N to darii, in waters offOregon. Fish. Bull., U.S. 75:173-183. 36”30’N). Southwest Fish Cent., Natl. Mar. Fish. Serv., NOAA, Pikitch, E., and R. Demory. 1988. Assessment of scales as a means of Tech. Memo. TM-NMFS-SWFC-133,44 pp. aging Dover sole. Trans. Am. Fish. Soc. 117:345-349. Childress, J. J., and M. H. Nygaard. 1973. The chemical composition Puckett, H. M. 1989. Ecology and possible causes of the jellied condi- of midwater fishes as a function of depth of occurrence off southern tion in Dover sole (Micvostomuspacificus).M. S. thesis, Humboldt State California. Deep-sea Res. 20:1093-1109. Univ., Arcata, Calif., 80 pp. Chilton, D. E., and R. J. Beamish. 1982. Age determination methods Quirollo, L. E, and P. Kalvass. 1987. Results of Dover sole tagging in for fishes studied by the groundfish program at the Pacific Biological waters off northern California, 1969-1971. Calif. Dep. Fish Game Station. Can. Spec. Publ. Fish. Aquat. Sci. 60, 102 pp. Mar. Res. Admin. Rep. No. 87-4, 73 pp. Demory, R. L. 1972. Scales as a means ofaging Dover sole (Micvosfomur Rofc D. A. 1982. Reproductive strategies in flatfish: a first synthesis. parifeus).J. Fish. Res. Board. Can. 29:1647-1650. Can. J. Fish. Aquat. Sci. 39:1686-1698. Dobush, G. R., C. D. Ankney, and D. G. Krementz. 1985. The effect -. 1983. An allocation model of growth and reproduction in fish. of apparatus, extraction time, and solvent type on lipid extractions of Can. J. Fish. Aquat. Sci. 40:1395-1404. snow geese. Can. J. Zool. 63:1917-1920. Sokal, R. R., and F. J. Rohlf. 1981. Biometry. 2nd ed. W. H. Freeman Draper, N., and H. Smith. 1981. Applied regression analysis. New arid Co., 859 pp. York:John Wiley & Sons, 709 pp. Somero, G. N., J. F. Siebenaller, and P. W. Hochachka. 1983. Physiolog- Engelman, L. 1988. LR stepwise logistic regression, In BMDP statisti- ical adaptations of deep-sea animals. III The sea, vol. 8: deep-sea cal software manual, vol. 2, W. J. Dixon, ed. Pp. 941-969. biology, G. T. Rowe, ed. N.Y.: Wiley Interscience. Eschmeyer, W. N., E. S. Herald, and H. Hammanti. 1983. A field guide Steven, G. A. 1930. Bottom fauna and the food of fishes. J. Mar. Biol. to Pacific Coast fishes of North America. Boston: Houghton-Mifflin U.K. 16677-706. Co., 336 pp. Templeman, W., and G. L. Andrews. 1956. Jellied condition in the Fisher, R. A,, R. A. Fritzche, and G. L. Hendrickson. 1987. Histology American plaice HippogIossoides platcssoides (Fabricius). J. Fish. Res. and ultrastructure of the ‘jellied’ condition in Dover sole, Micro- Bd. Can. 13:147-182. stomus pacifictts. Proc. V Congr. Europ. Ichthyol., Stockholm 1985, Wakefield, W. W. 1984. Feeding relationships within assemblages of pp. 345-350. nearshore and mid-continental shelf benthic fishes off Oregon. M. S. Gabriel, W. L., and W. G. Pearcy. 1981. Feeding selectivity of Dover thesis, Oregon State Univ., Corvallis, Oregon, 102 pp. sole, Micvosfomrrs pacificus, off Oregon. Fish. Bull., U.S. 79(4): Westrheim, S. J., and A. R. Morgan. 1963. Results from tagging a 749-763. spawning stock of Dover sole, Microsrornits pacificits. Pac. Mar. Fish. Hagerman, F. B. 1952. The biology of the Dover sole (Micvostomus Comm. Bull. 6:14-21. parifeus) (Lockington). Calif. Dep. Fish Game Fish Bull. 85, 48 pp. Yoklavich, M. M., and E. K. Pikitch. 1989. Fecundity and status of Harry, G. Y 1959. Time of spawning, length at maturity, and fecundity maturity of Dover sole, hfitrosrotnrts pacifetrs, off northern Oregon, of the English, petrale and Dover soles (Pavophvys t~etulirs,Eopsetta Fish Bull., U.S. 87(4). .jordani and Micvostomtrs patifcur, respectively). Fish. Comm. Oregon. Zar, J. H. 1984. Biostatistical analysis. New Jersey: Preiitice-Hall, Res. Briefs 7(1):5-13. 718 pp.

144 SMITH: MONITORING OF PACIFIC SARDINE SPAWNING AREA CalCOFl Rep., Vol. 31,1990

MONITORING INTERANNUAL CHANGES IN SPAWNING AREA OF PACIFIC SARDINE (SARDINOPS SAGAX) PAULE SMITH National Marine Fisheries Service Southwest Fisheries Center PO Box271 La Jolla, California 92038

ABSTRACT to 115,000 MT in the 1958-59 season; thereafter the It is easier to monitor the spawning area of the fishery declined to very low levels. Pacific sardine than to mount a full-scale effort to There appears to be some recovery of the Pacific precisely estimate spawning biomass. Monitoring the sardine stock (Parrish et al. 1989) off southern Cali- spawning area may be particularly economical when fornia (Wolf and Smith 1986; Wolf et al. 1987), as the sardine is extremely rare or extremely abundant. evidenced by the tendency for the spawning area to Such imprecise estimates will probably not answer increase. Since 1986, small quotas for commercial the management question of whether or not to set a catch have been permitted (907 MT for all use, 317 specific biomass quota. The spawning area estimate MT for live bait, 227 MT for dead bait). The inci- is a candidate - with aerial surveys, scale sedimen- dental catch of sardine has also increased, particu- tation rates, and acoustic-trawl surveys -for use in larly in the Pacific mackerel fishery (Wolf 1989). If interpolating between years when the more precise the spawning biomass continues to increase, there SWFC daily egg production method is used. will soon be a need for a management plan, and decisions will have to be made about monitoring the size of the stock. The moratorium on sardine catch RESUMEN for all uses was managed informally from 1974 to Resulta mis ficil monitorear el irea de desove de 1984 with an annual statement that the biomass ap- la sardina del Pacific0 que montar un estudio a gran peared to be below 20,000 short tons. Currently, the escala que estime con precisi6n la biomasa del de- spawning biomass of sardine is estimated from a sove. El monitoreo del irea del desove parece ser una spawning area relationship published by Wolf and salida economics cuando la abundancia de la sardina Smith (1985). The quota of 907 MT for all uses has es extremadamente baja o extremadamente alta. Es- been constant since 1986, but may be increased by tas estimaciones imprecisas probablemente no con- the California Department of Fish and Game. testen las preguntas administrativas relacionadas The Southwest Fisheries Center Coastal Division con la posibilidad de imponer una cuota a la biomasa has devised an absolute, instantaneous daily egg especifica. La estimacion del irea de desove junto production method to estimate spawning biomass con estudios akreos, determinaciones de velocidad of northern anchovy (Lasker 1985). It remains for a de sedimentacibn de las escamas, y estudios de a- management plan to determine how often and how rrastres actisticos sea probablemente el mejor mCtodo precisely the spawning biomass of sardine must be para interpolar datos entre aquellos aiios cuando el monitored. Annual management advice is currently mCtodo mas precis0 de la producci6n de huevos del based on the stock synthesis model (Methot 1989; SWFC es utilizado. Lo and Methot 1989;Jacobson and Lo 1989). Zweifel (1973) noted that the number of sardine INTRODUCTION eggs per positive station remained stable from 1951 The sardine fishery has been thoroughly reviewed to 1960, even as the population of sardine declined by Ahlstrom and Radovich (1970). The fishery be- by nearly an order of magnitude. Smith and Rich- gan in Monterey in the nineteenth century and grew ardson (1977: table 3.10) showed that the mean num- to a maximum seasonal catch ofjust over 700,000 ber of eggs per positive station varied little, even metric tons (MT) in the 1936-37 season. The catches with different quantitative net tows and population began to disappear from the Pacific Northwest and sizes varying from 4 million to 200,000 tons. Wolf northern California in 1945-46 and were inconse- et al. (1987: table 2) demonstrated that the mean quential by 1952-53. There was a minor resurgence number per positive net tow remained stable, with a sardine spawning biomass estimated at 20,000

~~ ~ ~~ [Manuscript received February 15,1990.1 tons; thus, over a measured range of 20,000 to 4

145 SMITH: MONITORING OF PACIFIC SARDINE SPAWNING AREA CalCOFl Rep., Vol. 31,1990

Figure 1. Locations of net tows taken to search for Pacific sardine eggs in April, May, and June 1929; April, May, and June 1930; February-August 1931; and February, April, and May 1932 (Scofield 1934). Solid circles indicate that the net tow was positive for eggs; open circles indicate that a net tow was taken and examined for eggs but none were found. million tons of spawning biomass, the distribution increase, and where further catches would probably of sardine eggs per positive station remains the delay the recovery of the stock to higher productiv- same. We believe that this circumstance arises from ity. The upper threshold is established at the point the schooling habit of adults, regardless of the where fleet and processor capitalization and costs of spawning biomass of the population; the further be- marketing would not be repaid owing to the tem- havioral concentration of sperm and eggs by the porary nature of high biomass. This requires rather fraction of fish involved in spawning at the time of precise monitoring of spawning biomass near the external fertilization in the open sea; and the subse- lower threshold, and less precise estimates at all quent dispersal by turbulent diffusion (Smith 1973; other biomass levels. Also, the policy of using in- Smith and Hewitt 1985; Mangel 1985; Mangel and dices of abundance rather than annual absolute mea- Smith, in press). Theoretically, the number of eggs sures of abundance reduces management costs for per positive station could decline at high biomass assessing stock at higher levels of abundance while concentrations because filter feeding in schooling providing for a stable fishery. fishes like the sardine and anchovy may result in It is the purpose of this paper to present data to incidental cannibalism and predation of the eggs aid in designing programs to estimate and monitor (Gulland 1971; Alheit 1987; Smith et al. 1990). biomass. These programs should contribute to It is assumed that the management plan for sar- management of the Pacific sardine fishery and to the dine will be similar to that for anchovy, with two understanding of interactions among the sardines, thresholds (PFMC 1983). When the spawning bio- other planktivorous fishes, and their environment. mass is below the lower threshold, no fishery is per- The data used in the original paper on this topic by mitted; when the spawning biomass is between the Zweifel (1973) will be extended from 1940 to the lower and upper threshold, a fixed fraction of the present. spawning biomass, based on demographic consid- erations, is established as a quota for the ensuing year; and when the spawning biomass is above the METHODS AND RESULTS upper threshold, no further catch is authorized. The Surveys of sardine eggs and larvae were con- lower threshold is established at the point that the ducted in 1929-32 (Scofield 1934; figure l), 1939- costs of the fisher’s search for the remaining fish 1941 (Ahlstrom 1948; Ahlstrom 1966; Smith 1972;

146 SMITH: MONITORING OF PACIFIC SARDINE SPAWNING AREA CalCOFl Rep., Vol. 31,1990

53Columbia River t 0 0 00 “I

1;Cape Mendocino

Figure 3. Contoured abundance of sardine eggs during the period of survey in 1941 (Sette and Ahlstrom 1948). These data are included in the analysis.

contains all the biomass, is a consistent fraction of 0 0. the total spawning area, or contains a constant frac- 0. tion of the spawning biomass. It is likely that all time 0 series used will suffer from violation of the same 0 assumptions, since all. values contain measurement error, and the values are drawn from an autocorre- 0 lated time series. I do not believe the regression 0 methods used (Williams 1983) are valid for direct 0 estimates of spawning biomass for management purposes. Recognizing these limitations, I will use the well-known procedures for linear regression to evaluate the components of indirect measures of 0 population size. 1939 Sardines 0 0 Regress ion Analysis o No eggs Data on egg and larval abundance are available for 0 Eggs present 0 0O0 0 selected years for a major region of spawning. In a I 0 previous analysis of anchovy and sardine (Smith 1972), only the larval data were used. It was noted Figure 2. Locations of net tows taken to search for Pacific sardine eggs in May that larvae can be sampled over 3 weeks and disperse and June 1939 (Ahlstrom 1948). Net tows taken by Scofield (1934) were used and cover more area than eggs, which can only be to design the cruise. sampled over 3 days. In addition, the eggs retain the distributional characteristics of the schooled adults figure 2); and 1951-89 (CalCOFI on-line data sys- that spawned them. Although larvae have the statis- tem). Although distributions of eggs and larvae per tical advantage of dispersal, over 3 weeks’ time there net tow can be obtained for all these time periods, it may be considerably more variability in mortality is not possible to measure the areal boundaries of the than for eggs. In this study I use both egg and larval spawning distribution for all sets of years. For this data as well as the biomass estimates from virtual illustration, I have chosen the time series of sardine population methods used for catch analysis from eggs and larvae for the area surveyed in 1941 (figure 1932 through 1965 (Murphy 1966: 1932-44; MacCall 3). At that time, virtual population estimates were 1979: 1945-65; table 1). For the remainder of the text based on the assumption that every female over 2 I use the notation as follows: years old was a spawner; maturity and gonadal ac- tivity were not checked in each year. VPM- the biomass (MT) of sardines 2 years old The method used here for estimating biomass is and older; the “index area” method. The quality of this method E- the egg census estimate of abundance in the depends on the assumptions that the area chosen entire survey area (#/lorn’);

147 SMITH: MONITORING OF PACIFIC SARDINE SPAWNING AREA CalCOFl Rep., Vol. 31,1990

TABLE 1 Omission of the constant when evaluating PE did Data for Regression Estimates Using the Census Estimate not materially change the F value of the analysis of or the Partitioned Estimates of Eggs or Larvae and the variance. Virtual Population Methods ( VPM) for Pacific Sardine Possibly the most significant result of this analysis Year N E L PE PL VPM for monitoring the sardine biomass is the weakness ~~~~~_____~~ ~~ ~~~ A. Years for which VPMis available of number of eggs or larvae per positive station 1940 240 699.10 49.13 0.754 0.808 1759.6 when coupled with the probability of a positive sta- 1941 210 336.90 36.88 0.629 0.748 2457.1 1951 96 33.33 2.06 0.167 0.146 277.0 tion; in both cases p is greater than .05 and negative 1952 152 6.85 3.49 0.099 0.105 136.0 in reflecting changes in biomass (PE and NE, PL and 1953 226 0.21 0.07 0.031 0.013 202.0 NL in table 2; see also NE in figure 4B). As in the 1954 219 39.67 11.98 0.146 0.110 239.0 1955 142 26.87 7.29 0.169 0.092 170.0 previous analysis (Smith 1972), the constants are not 1956 156 47.88 6.90 0.090 0.045 108.0 important, and the larval time series, with or with- 1957 145 23.00 12.09 0.103 0.097 90.0 out inclusion of number per positive station (NL), 1958 171 86.38 8.86 0.298 0.310 177.0 1959 188 182.00 7.88 0.287 0.250 122.0 are marginally better predictors of spawning bio- 1960 197 117.14 4.47 0.183 0.168 88.0 mass (actually biomass of age 2 + ) than the equiva- 1961 73 17.08 1.40 0.164 0.082 54.0 lent measures of eggs (table 2). This may simply 1962 64 1.77 0.95 0.016 0.047 27.0 1963 77 14.22 1.94 0.052 0.039 21.0 reflect the longer duration and better mixing with 1964 183 0.43 0.00 0.022 0.005 11.0 resultant lower variance of the larval estimates. I use 1965 112 4.57 0.79 0.107 0.098 3.0 the proportion of eggs relationship (PE) to extend B. Years for which no VPM is available 1966 169 2.01 0.29 0.012 0.053 the biomass time series to the present (table 3;figure 1969 147 0.33 0.26 0.027 0.041 4A). 1972 118 0.00 0.03 0.000 0.008 Another significant result is that the standard er- 1975 267 2.54 0.07 0.026 0.007 1978 189 0.38 0.18 0.026 0.016 rors of the parameter estimates are all relatively 1981 139 0.99 0.23 0.029 0.007 large. This implies than none of these estimates 1984 141 3.40 6.50 0.064 0.043 would be adequate for year-by-year management 1985 99 10.96 8.80 0.061 0.051 1986 I83 3.45 3.62 0.011 0.044 advice during close regulation of the fishery. It 1987 81 18.73 23.00 0.062 0.111 seems likely that some of this parameter error is due 1988 85 40.75 2.25 0.082 0.047 to the fixed maturation at 2 years of age. The largest 1989 72 61.00 4.96 0.111 0.167 ~ ~~ ~~ ~~~ ~ ~~~~ deviations occur at the times of significant temper- Key: N = number of net tows included, E = egg census estimate of abundance; L = larval census estimate; PE = proportion of net tows ature anomalies. It may be that temperature plays an containing sardine eggs; PL = proportion of net tows containing important role in determining the rate of maturation sardine larvae; VPM = biomass of sardines 2 years old and older. in sardine, as has been found for anchovy (Methot 1989). If the age at first maturity is shown to be L - the larval census estimate (#/lorn’; Smith influenced by temperature, it is likely that the VPA 1972); series could be adjusted from temperature records PE- the proportion of the net tows containing on hand (see Methot 1989, table 1). sardine eggs (positive stations); Even with the improvement in accuracy, it does PL - the proportion of the net tows containing not appear from table 3 that useful estimates can be sardine larvae; obtained from the level of sampling effort now being NE- the number of eggs per positive station (#/ conducted in the quarterly CalCOFI surveys. Al- 10m’); and though the trend of recovery may be essentially cor- NL-the number of larvae per positive station rect, the spawning biomasses indicated may well be (#/lorn”). overestimates (Wolf and Smith 1986; Wolf et al. 1987). In the absence of an adequate SWFC daily egg NE and NL are not listed in table 1 because they are production method estimate to provide a recent cal- obtained simply by dividing the census estimate (E) ibration, it does not seem likely that spawning area by the proportion of positive stations (PE). estimates alone will be adequate for setting quotas for the Pacific sardine fishery (figure 4A). The Table 2 lists the parameters, the standard errors of spawning area estimate would be useful for moni- estimate of the parameters, the Student’s t value of toring the sardine stock at high levels, and the area the estimate, the probability of the t value, the F and VPM or stock synthesis estimates may be useful value of analysis of variance, and the probability of for examining this population’s impact on the eco- the F value. No constant is less than .05, thus all the system. If the stock recovers to more than a million equations were redone forcing the zero-intercept. tons it may be necessary to conduct wide-ranging

148 SMITH: MONITORING OF PACIFIC SARDINE SPAWNING AREA CalCOFl Rep., Vol. 31,1990

TABLE 2 Comparison and Evaluation of Regression Parameters for Indirect Estimation of Spawning Biomass of Pacific Sardine

Predictor Coef SD Student’s t V F-ratio ~ ~~~~ P A. With constani Constant 51.2 113.5 0.45 0.659 29.05 0.000 E 3.0976 0.5747 5.39 0.000

Constant -68.40 88.15 -0.78 0.450 68.12 0.000 L 45.489 5.511 8.25 0.000

Constant -225.5 108.7 - 2.07 0.056 57.00 0.000 PE 2946.9 390.3 7.55 0.000

Constant - 143.87 83.06 -1.73 0.104 88.96 0.000 PL 2651.7 281.1 9.43 0.000

Constant -77.5 214.0 -0.41 0.722 6.98 0.018 NE 1.4407 0.5453 2.64 0.018

Constant 296.4 252.3 1.17 0.258 0.08 0.780 NL 1.114 3.921 0.28 0.780

Constant - 85.78 91.08 -0.94 0.362 33.89 0.000 E - 1.034 1.188 -0.87 0.399 L 58.22 15.64 3.72 0.002

Constant - 158.8 115.2 -1.38 0.190 31.42 0.000 PE 3559.9 574.2 6.20 0.000 NE -0.6284 0.4432 -1.42 0.178

Constant - 175.0 111.4 -1.57 0.139 42.17 0.000 PL 2647.8 289.2 9.16 0.000 NL 0.669 1.536 0.44 0.670

Constant -67.0 113.2 -0.59 0.565 28.39 0.000 PE - 2425 2019 -1.20 0.253 PL 5214 1669 3.12 0.009 NE -0.6225 0.3882 - 1.60 0.135 NL 2.183 1.519 1.44 0.176 B. No constant E 3.2239 0.4891 6.59 0.000 43.44 0.000

L 43.032 4.455 9.66 0.000 93.30 0.000

DE 2379.4 305.9 7.78 0.000 60.52 0.000

PL 2345.1 231.6 10.12 0.000 102.50 0.000

NE 1.2915 0.3476 3.72 0.002 13.80 0.002

NL 4.527 2.663 1.70 0.109 2.89 0.109

E - 0.789 1.154 -0.68 0.505 45.33 0.000 L 52.27 14.26 3.67 0.002

PE 3469.8 587.3 5.91 0.000 38.98 0.000 NE - 0.8776 0.4166 -2.11 0.052

PL 2437.8 268.7 9.07 0.000 49.93 0.000 NL -0.881 1.234 -0.71 0.486

PE - 3963 1758 -1.69 0.116 39.12 0.000 PL 5620 1484 3.79 0.002 NE -0.6318 0.3781 - 1.67 0.119 NL 1.832 1.364 1.34 0.202

Key: E = egg census estimate ofabundance; L = larval census estimate; PE = proportion ofnet tows containing sardine eggs; NE = niiniber of eggs per positive station; PL = proportion ofnet tows containing sardine larvae; NL = number of larvae per positive station.

149 SMITH: MONITORING OF PACIFIC SARDINE SPAWNING AREA CalCOFl Rep., Vol. 31,1990

- 1,000 -h 900 6 2 800 .--0 = VPM 8 700 hg A = VPMHAT v) :E 600 0: UU' 500 Z$ U 400 uw In0 2 300 U 3 200 5 = 100

0 0 30 40 50 60 70 80 90 40 50 60 70 80 90 A YEAR B YEAR

Figure 4. Time series plot of sardine 1932-89 (Murphy 1966: 1932-44; MacCall 1979: 1945-65). A, estimates using virtual population methods or egg incidence correlates (PO;B, estimated egg abundance per unlt surface area at positive stations (NE).

TABLE 3 surveys (figures 1 and 2) to accomplish a daily egg Virtual Population Estimates of Sardine Spawning production estimate based on the extent of spawn- Biomass (2 + ) Derived from Regression on the ing in the 1929-32 and 1939 surveys (Scofield 1934; Proportion of Positive Egg Stations in 1940-41 Ahlstrom 1948). and 1951-65 Year N PE VPMHAT ~ ~~ ~ ~~~~ DISCUSSION 1940 240 0 754 1800 Monitoring sardine biomass for fisheries manage- 1941 210 0 629 1500 1951 96 0 167 400 ment and ecosystem time series can be done with 1952 153 0 099 240 precise, but elaborate and expensive, egg production 1953 226 0 031 74 methods as used for the baseline studies of northern 1954 219 0 146 350 1955 142 0 169 400 anchovy (Lasker 1985). Alternatively, indirect meth- 1956 156 0 090 210 ods could be used, such as the deposition rate of 1957 145 0 103 250 scales (Soutar and Isaacs 1974) in anoxic sediments 1958 171 0 298 710 1959 188 0 287 680 or traps; aerial surveys (Squire 1972); incidence and 1960 197 0 183 440 abundance of eggs or larvae; or trawl-acoustic meth- 1961 73 0 164 390 ods in a mixed technique model (Methot 1989). Sar- 1962 64 0 016 38 1963 77 0 052 120 dine stocks in the California Current region and 1964 183 0 022 52 other regions in the world fluctuate by orders of 1965 112 0 107 250 magnitude (Smith and Moser 1988; Lluch-Belda et 1966 169 0 012 29 1969 147 0 027 64 al. 1989), and rough estimates of biomass may be 1972 118 0 000 0 sufficient for management plans during many pe- 1975 267 0 026 62 riods of the species time sequence. 1978 189 0 026 62 1981 139 0 029 69 1984 141 0 064 150 ACKNOWLEDGMENTS 1985 99 0 061 150 I thank Patty Wolf, John Hunter, Izadore Barrett, 1986 183 0 011 26 1987 81 0 062 150 Marc Mangel, Alec MacCall, and an unnamed re- 1988 85 0 082 200 viewer for offering suggestions for materially im- 1989 72 0 111 260 ~ ~ ~ ~~ ___ proving the text. I am particularly grateful to Key N = number ofnet tows, PE = proportion ofnet tows Richard Charter and Cindy Meyer for maintaining containing sardine eggs, VPMHAT = estimate of sardine biomass 2 years old and older based on the regression estimate of table 2, no the CalCOFI data base and writing special programs constant for extracting the data.

150 SMITH: MONITORING OF PACIFIC SARDINE SPAWNING AREA CalCOFl Rep., Vol. 31,1990

LITERATURE CITED PFMC: Pacific Fishery Management Council. 1983. Northern anchovy Ahlstrom, E. H. 1948. A record of pilchard eggs and larvae collected fishery management plan. PFMC, 526 S. W. Mill St., Portland, Ore- during surveys made in 1939 to 1941. Spec. Sci. Rep. U.S. Fish and gon 97201. Wildl. Serv. Fisheries (54), 82 pp. Scofield, E. C. 1934. Early life history of the California sardine (Sardiria -. 1966. Distribution and abundance of sardine and anchovy lar- raerrilea), with special reference to distribution of eggs and larvae. vae in the California Current region off California and Baja Califor- Calif. Dep. Fish Game Fish Bull. 41:l-48. nia, 1951-64, a summary. Spec. Sci. Rep. U.S. Fish and Wildl. Serv. Sette, 0.E., and E. H. Ahlstrom. 1948. Estimations of abundance of Fisheries (534), 71 pp. the eggs of Pacific pilchard (Sarditiops raenrlra) off southern California Ahlstrom, E. H., and J. Radovich. 1970. Management of the Pacific during 1940 and 1941. J. Mar. Res. 7:511-542. sardine. It1 A century of fisheries in North America, N. G. Benson, Smith, P. E. 1972. The increase in spawning biomass of northern an- ed. Special Publication No. 7, American Fisheries Society, Wash. chovy, Brgvaulis morda.~.Fish. Bull. U. S. 70:849-874. D.C., pp. 183-193. -. 1973. The mortality aiid dispersal of sardine eggs and larvae. Alheit, J. 1987. Cannibalism versus egg predation: their significance in Rapp. P. -V. Reun. Cons. Int. Explor. Mer 164282-292. anchovies. In The Benguela and comparable ecosystems, A. I. L. Smith, €? E., and R. l? Hewitt. 1985. Anchovy egg dispersal and mor- Payne, J. A. Gulland, and K. H. Brink, eds. S. Afr. J. Mar. Sci. 5:467- tality as inferred from close-interval observations. Calif. Coop. 470. Oceanic Fish. Invest. Rep. 26:97-110. Gulland, J. A. 1971. Ecological aspects of fishery research. lii Advances Smith, P. E., and H. G. Moser. 1988. CalCOFI time series: an overview in ecological research, volume 7, J. B. Cragg, ed. New York: Aca- of fishes. Calif. Coop. Oceanic Fish. Invest. Rep. 2966-78. demic Press, pp. 115-176. Smith, €? E., and S. L. Richardson. 1977. Standard techniques for pe- Jacobson, L. D., and N. C. H. Lo. 1989. Spawning biomass of the lagic fish egg and larva surveys. FA0 Fisheries Tech. Pap. 175, FA0 northern anchovy in 1989. Southwest Fisheries Center Administra- Rome, 100 pp, tive Report LJ-89-17. Smith, P. E., H. Santander, atid J. Alheit. 1990. Comparison of mortal- Lasker, R., ed. 1985. An egg production method for estimating spawn- ity rates of Pacific sardine, Smdiiiops sagax, and Peruvian anchovy, ing biomass of pelagic fish: application to the northern anchovy, Eli- Etip~~lirr;iigem, eggs off Peru. Fish. Bull. U. S. 87:497-508. gradis mordax. NOAA Tech. Rep. NMFS 36, 99 pp. Soutar, A,, aiid J. D. Isaacs. 1974. Abundance of pelagic fish during the Lluch-Belda, D., R. J. M. Crawford, T. Kawasaki, A. D. MacCall, 19th and 20th centuries as recorded in anaerobic sediment off the R. H. Parrish, R. A. Schwartzlose, and P. E. Smith. 1989. Worldwide Californias. Fish. Bull. U. S. 72257-273. fluctuations of sardine and anchovy stocks: the regime problem. Squire, J. L., Jr. 1972. Apparent abundance ofsome pelagic marine fishes S. Afr. J. Mar. Sci. 8:195-205. off the southern and central California coast as surveyed by an air- Lo, N. C. H., and R. D. Methot. 1989. Spawning bioniass of the borne monitoring program..Fish. Bull. U.S. 70:1005-1019. northern anchovy in 1988. Calif. Coop. Oceanic Fish. Invest. Rep. Williams, B. K. 1983. Some observations 011 the use of discriminant 30:18-31. analysis in ecology. Ecology 64:1283-1291. MacCall, A. D. 1979. Population estimates for the waning years of the Wolf, I! 1989. Review ofsome California fisheries for 1988. Calif Coop. Pacific sardine fishery. Calif Coop. Oceanic Fish. Invest. Rep. 20:72- Oceanic Fish. Invest. Rep. 30:7-17. 82. Wolf, P., and P. E. Smith. 1985. An inverse egg production method for Mangel, M. 1985. Search models in fisheries and agriculture. Lect. determining the relative magnitude of Pacific sardine spawning bio- Notes Biomath. 61:105-138. mass off California, Calif. Coop. Oceanic Fish. Invest. Rep. 26:130- Mangel, M., and P. E. Smith. In press. Presence-absence sampling for 138. fisheries management. Can. J. Fish. Aquat. Sci. 47. . 1986. The relative magnitude ofthe 1985 Pacific sardine spawn- Methot, R. D. 1989. Synthetic estimates of historical abundance and ing biomass off southern California. Calif. Coop. Oceanic Fish. In- mortality for northern anchovy. Iii Mathematical analysis of fish vest. Rep. 272-31. stock dynamics, E. F. Edwards and B. A. Megrey, eds. Am. Fish. Wolf, P., €? E. Smith, and C. L. Scannell. 1987. The relative magnitude SOC.Symp. 6:66-82. of the 1986 Pacific sardine spawning biomass off California. Calif Murphy, G. I. 1966. Population biology ofthe Pacific sardine (Surdimps Coop. Oceanic Fish. Invest. Rep. 282-26. raerrrlea). Proc. Calif. Acad. Sci. Fourth Series 34:(1)1-84. Zweifel, J. R. 1973. A tion-parametric approach to the estimation of Parrish, R. H., R. Serra, and W. S. Grant. 1989. Monotypic sardines, relative change in fish population size from egg and larval surveys. Sardim and Snrdiriops: their taxonomy, distribution, stock structure, Rapp. P. -V. Reun. Cons. Int. Explor. Mer 164276-281. and zoogeography. Can. J. Fish. Aquat. Sci. 46:2019-2036.

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CalCOFI REPORTS -INSTRUCTIONS TO AUTHORS

Manuscvipt should be typed (no dot-matrix printouts, please) (chapter): double-spaced with wide, ragged right margins, and submit- Wooster, W. S., and J. L. Reid, Jr. 1963. Eastern boundary currents. In ted complete with figures, figure captions, and tables, in tripli- The sed, M. N. Hill, ed. New York: Intersclence Pub. pp. 253-280. Tables (with arabic numbers) should be typed (double- cate to spaced) separately from the text; each table should start on a CalCOFI Coordinator separate page and must have a brief title. Please avoid vertical California Department of Fish and Game rules and be consistent in format. 330 Golden Shore, Suite 50 Figures, whether drawings or halftones, should be submitted Long Beach, CA 90802 in a format not larger than 8% x 11”. Submit one set of camera-ready figures plus 2 sets of copies. Photographs should Manuscripts must be received by February 1 of the year in be printed on glossy paper. Drawings should be reduced pho- which publication is desired. tographically. A composite figure should be submitted as a Sequence of the material should be TITLE PAGE, AB- single photograph or at least as a single careful paste-up. Figures STRACT, RESUMEN, TEXT, LITERATURE CITED, will appear as either single-column (85-mm-width limit), or APPENDIX (if any), FOOTNOTES (if any), TABLES, LIST double-column (178-mm-width limit); or as full page. Special OF FIGURES with entire captions, and FIGURES. cases should be discussed with the editor before submittal. After reduction, no letter or number should be smaller than 1 Title page should give: mm. Special note should be taken of the disappearance of deci- a running head of no more than 60 letters and spaces mal points during reduction. If commercially prepared shading title of the article is used, make a trial reduction to ensure that the patterns do not author(s) name(s) and affiliation(s) merge at the required reductions. The determining factor for address(es), including Zip Code(s) size should be the complexity of detail to be shown. Abstract should not exceed one double-spaced page and Each figure must have a caption; captions should be typed, must be submitted both in English and in Spanish (Resumen). double-spaced, in numbered sequence on a separate sheet. Text style will in general follow that of the U. S. Department Illustrative materials submitted for publication are often first of Commerce (NOAA) Fishery Bulletin. Contributors who are prepared for oral presentation in slide format. Authors should not familiar with this publication will do well to follow The take special care that slide-format material submitted to Cal- Chicago Manual of Style (1982). Authors are strongly urged to COFI Reports is appropriate to printed format with respect to compare their typewritten equations with similar expressions economy, redundancy, and style. in the printed literature, with special attention to ambiguity of Acknowledgments, if included, should be placed at the end of the symbols for “one” and for “el,” before submitting. When- the text and may include funding source. ever possible, write in the first person, and use active verbs. Repvint orders will be mailed (to senior author only) on pub- Measurements must be given in metric units; other equivalent lication of the completed book. No covers will be supplied, and units may be given in parentheses. there will be no further reproduction. Personal communications and unpublished data should not be The CalCOFI Reports will use the CalCOFI Atlas full-page included in the Literature Cited but may be cited in the text in chart format where the material would be best used overlaid on parentheses. Use footnotes only when parentheses will not suf- the CalCOFI Atlas charts for purposes of comparison and fice. List footnotes on a separate sheet. where the material presented is of insufficient scope and quan- Literature.cited should appear in the text as Smith (1972) or tity to warrant the publication of an atlas. Smith and Jones (1972) or (Smith and Jones 1972; Jones and The CalCOFI Editorial Board will consider for publication, Smith 1973) or Smith et al. (1972). All literature referred to in in the section entitled “Scientific Contributions,” manuscripts the text should be listed (double-spaced) alphabetically by the not previously published elsewhere that bear some relationship first author on a separate sheet under the heading Literature to the following with respect to the Californias, the California Cited. Only the authors’ surnames and initials will be used. No Current, and the Gulf of California: citation should appear in the list of Literature Cited unless it is marine organisms cited in the text, tables, or figure captions. Each citation must marine chemistry, fertility, and food chains be complete according to the following: marine fishery modeling, prediction, policy, and manage- (article): ment Eppley, R. W., E. H. Renger, E. L. Venrick, andM. M. Mullin. 1973. marine climatology, paleoclimarology, ecology, and paleo- A study of plankton dynamics and nutrient cycling in the central ecology gyre of the North Pacific Ocean. Limnol. Oceanogr. 18(4):543-551. (book): marine pollution Odum, E. F’. 1959. Fundamentals of ecology. 2d ed. Philadelphia: physical, chemical, and biological oceanography Saunders, 546 pp. new marine instrumentation and methods.

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CALCOFI BASIC STATION PLAN -. SINCE 1950 CalCOFl Rep., Vol. 31,1990

CONTENTS

In Memoriam ...... ,...... 5 I. Reports, Review, and Publications Report of the CalCOFI Committee ...... 7 Review of Some California Fisheries for 1989 ...... 9 Publications ...... 22 11. Fortieth Anniversary Symposium of the CalCOFI Conference, 1989 OCEAN OUTLOOK: GLOBAL CHANGE AND THE MARINE ENVIRONMENT ...... 25 Presentation to Roger Revelle ...... 28 Remarks by the Honorable Leo McCarthy, Lieutenant Governor of California ...... 30 Remarks by the Honorable John Knauss, Under Secretary of Commerce for Oceans and Atmosphere ...... 33 Panel Discussion: What Society Needs from Ocean Scientists in Preparing for Global Change. Robert Sulnick, Byvon Shev, Edward Fvieman, Bert Lavkins, John McGowan, Boyce Thovne Millev, and Havvy Scheibev ...... _...... _...... __...... _.... 37 111. Scientific Contributions California Marine Research and the Founding of Modern Fisheries Oceanography: CalCOFI’s Early Years, 1947-1964. Harry N. Scheibev ...... 63 Settlement ofJuvenile California Halibut, Pavalichthys cal$ovnicus, along the Coasts of Los Angeles, Orange, and San Diego Counties in 1989. M.James Allen and Kevin T Hevbinson . . . 84 Genetic Structure of White Seabass Populations from the Southern California Bight Region: Applications to Hatchery Enhancement. Devin M. Bavtley and Donald B. Kent ...... 97 The Vertical Distribution and Feeding Habits of Two Common Midwater Fishes (Leuvoglossus stilbius and Stenobrachius leucopsavus)off Santa Barbara. Gvegov M. Cailliet and Alfved W Ebeling ...... , ...... 106 The Market for Fish Meal and Oil in the United States: 1960-1988 and Future Prospects. Cynthia J. Thomson ...... 124 Bathymetric Patterns in Size, Age, Sexual Maturity, Water Content, and Caloric Density of Dover Sole, Micvostomuspacijicus.John R. Hunteu,John L. Butlev, Cavol Kimbvell, and Eric A. Lynn ...... 132 Monitoring Interannual Changes in Spawning Area of Pacific Sardine (Savdinopssagux). Paul E. Smith ....._...... 145 Instructions to Authors ...... 152 CalCOFI Basic Station Plan ...... inside back cover

Indexed in Cuvvent Contents. Abstracted in Aquatic Sciences and Fisheries Abstracts and Oceanic Abstracts.